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BIOLOGICAL ORDER

KARL TAYLOR COMPTON
The Karl Taylor Compton
Technology by bringing
of the ninth president of the

1881-1954

Lectures honor the

memory

Massachusetts Institute of

to the M.I.T.

community some

of the great minds of our time

who

contribute to the inte-

gration of scientific, cultural, and philosophical concerns
a synthesis richly

achieved by Karl Taylor

Compton and
long leader-

shared with colleagues and students during
ship of the Institute.

his

.KARL TAYLOR COAIPTON LECTURERS Niels Bohr 1937 Otto Struve 1939 Andre Lwoff I960 Rabi 1962 Isidor I.

KARL TAYLOR COMPTON LECTURES .

BIOLOGICAL ORDER by Andre Lwoff The M-I-T Press Massachusetts Institute of Technology Cambridge Massachusetts .

62-16929 Printed in the United States of America .Copyright © 1962 The Massachusetts Institute of Technology All rights reserved Library of Congress Catalog Card' No.

as if the The being intended for physicists and chemists. despite the fact that intended for this institute enjoys an I excellent department of biology. I certainly have to apologize for this hypothesis. mostly physiAfter discussing this matter with my M. in its VVe have material. It is in view of this ambitious aim that biological order was selected as a theme. were planned audience knew nothing about cell biology. are cists The Compton Lectures. Thus life. Biological order has been considered at the molecular level in static as its well as in its dynamic aspects. I decided that these Compton Lectures would be directed toward the young physicists and chemists. struc- reproduction. namely the nature. I have tried to explain how [ vii ] . colleagues. and viruses representative of a specific order and at the same time of disorder. the organism. told.T. and the problem of biological order posed ture. dis- cussed in succession: the hereditary order. to interest them in biological problems.PREFACE A scientist invited to deliver a series of lectures has first to select a theme which is determined essentially by his own field of interest and activity. have been defined and the generality. namely the control of interaction enzyme synthesis. of and variation of the genetic the functional order.I. as was and chemists. a namely lectures. with a very specific goal. This is especially important when a biologist is going to lecture in an institute of technology. The scientist then has to organize the lectures according to the interest and knowledge of the potential audience. finally and the as heredity and environment. very large group of students.

will be with the is impression that biological order. and trols how the organism con- and adjusts I its molecular balance. 1960 [ viii ] . fascinating subject. a Andre Lwoff Paris May. after having penetrated left into the molecular intimacy of the living being.PREFACE molecules communicate and interact. the foundation of biology. May express the hope that the reader.

CONTENTS I. ' 4 5 Organism. THE PROBLEMS Organism and Molecules 7 7 The Subject Metabolism Simplicity or Complexity 9 9 10 10 Unity III. THE HEREDITARY ORDER: GENETIC INFORMATION Birth of the 14 14 15 17 Modern Concept Necessity of Variation The Genetic Material Identified Transformation Structure of the Genetic Material Desoxyribonucleic acid Ribonucleic acid 17 19 19 21 The Molecular Mutation Basis of Enzymatic Specificity 22 Protein Synthesis: The Code The nucleic acid The "mutated" protein ' 24 26 26 28 80141 [ix] . and Assimilation II. INTRODUCTION TO BIOLOGY Life 3 3 3 The Organism The Cell and Organisms Life. Reproduction.

inhibition of enzymatic activity The 54 58 58 58 Preclusive and corrective feedback mechanisms Conclusion 60 V. sexual digression 39 39 41 43 43 46 Altered enzymes Induction and Repression Generalized induction hypothesis The repressor The Ways of Repression Repressor. lysogeny 65 65 66 66 68 [X] . Amino acids 49 49 53 Carbon and energy source operator Co-ordinated control Refinement of regulation. THE FUNCTIONAL ORDER The Problem 35 35 Two Models of Enzyme Synthesis Induction 37 37 38 Repression The Genetic Control of Bacterial Metabolism Metabolism of lactose Variations The gene as a unit.Reproduction of the Genetic Material DNA Production of an Reproduction of Conclusions 29 29 32 RNA template from DNA RNA 32 33 IV. or virion The virus Remarks on the control of viral functions Bacteriophage Life-cycle of a virulent bacteriophage The temperate bacteriophage. VIRAL FUNCTIONS: ORDER DISORDER AND 62 62 63 Introduction Viral order: the infectious viral particle. aporepressor. corepressor. and the control of enzyme syntheses 46 46 49 inducer.

The prophage The receptor Prophage and vegetative phage 69 70 70 71 71 Immunity Induction Inducing agents Zygotic induction Infection and lysogenization Mutations affecting inducibility Mutations affecting virulence Mutations affecting the vegetative phase Interactions of Cellular and Viral Functions Cellular control of viral reproduction Viral control of cellular functions Viral and nonviral diseases Viruses and cancer 73 73 75 75 77 78 80 80 82 83 83 VI. BIOLOGICAL ORDER AND ENTROPY Sources of Energy Equations 87 87 88 89 90 91 Entropy Life and Entropy of Biological Organization Information and Negentropv The Negentropy Remarks on "Genetic Information" Entropy and Evolution 92 93 94 95 The Source of Orderliness VII. CONCLUSION 99 .

.

specific system able to reproduce its kind. organized. such does not has nobody the has ever seen being no such thing. or as a state of organisms. the study of easiest solution is life. this exercise a matter of an excellent to condense the essential of a phenomenon it into a formula — the formula conit taining everything has to contain. identical organism. as so people have done. is. . and excluding everything To cast a critical good definition is therefore useful." Yet. [3] . It is. Szent-Gyorgyi writes: "Life it. An organism never appears de organism An novo but always derives from a pre-existing. . Life may be considered either as a property. by etymology. In his as many book The exist. Life is and the its to decide. that definition is impossible. The physicist will immediately ask two questions: (1) What an organism? (2) What is the specific property^ of living organisms that does not exist in the inanimate world and is therefore is characteristic of life? The Organism may be visualized as a complex. as heuristic method. Nature of Life. among methods for discovery.INTRODUCTION TO BIOLOGY Life Biology difficult to define. For it obliges one category or of has to exclude. or as a mamfestatio7i. it is The noun 'life' well known that a no sense. for the terms or aspects compels consideration of all of a problem. This might or might not satisfy the biologist. there is definition fact. .

BIOLOGICAL ORDER Biological systems continuity. This is us consider. it shows dark central body. surrounded by a pale substance called cytoplasm. independent unit of integrated structures and functions that reproduces true to type can only be an organism. The cells of a multicellular organism are specialized. It is a sphere 1 in diameter. in his fascinating book Cybernetics: "Living organisms are metastable Maxwell demons whose stable state is to be dead. Let bacterium. And such a complex. among which are nucleic acids and proteins. composed essentially of macromolecular compounds. and rethe independent unit of reproduction. grows. When provided with food. and some would perhaps prefer a more original and sophisticated definition. Syntheses can take provided with food and energy. which is consumed in the processes involved. Reproduction of a complex system containing macromolecules is therefore characteristic of life. the group to which we are so proud to belong. this metabolism. Leydig described and defined the cell as a mass of protoplasm containing about a a nucleus. When properly handled and stained. a living organism. produces. that is. An The simplest organism is therefore a relatively complex machine. a microbial /i. cell. The formulation which follows is an attempt at a summary of the views expressed by Norbert Wiener." The Cell and Organisms Organisms belong to two categories: {a) the protists or microbes. the nucleus. Even the smallest organism contains a few thousand different species of organism macromolecules. for example. The liver cells are different from the kidney cells [4] . organisms — are endowed with genetic The organism place only are if synthesizes is the system its constitutive parts. (^) those which are composed of many cells. the is The organism is organism metabolizes. which are composed of one cell and are therefore the lower organisms. differentiated. a an excellent definition. In 1857. All know^n complex systems which contain macromolecules and are able to reproduce their kind belong to the living systems. — that is. the so-called higher organisms. These statements might be considered too factual.

"the action the assemblage of small building blocks of making like. share so As such. the microbial to a complete organism. co-operation. The functioning The cells of of each cell is a multicellular organism are the integrated. and interdependent parts of the organism. Assimilation essentially is the organization of complex specific sequences. into specific macromolecules for — proteins or nucleic acids — that are characteristic each species. Organism. [5] ." — amino acids or nucleic bases — common is to all living beings. Assimilation of life. assimilation. cell. contain Organisms ated. Assimilation a patternization. But cell essential to know word applies to two different categories of systems: the de- pendent part of the multicellular organism and the independent unicellular organism. In order that these can be perpetu- smallest unit of integration. the other. putting each of the twenty amino acids and each of the four nucleic bases where it belongs. Only a living being is able to s

nthesize a few thousand specific macromolecules. an independent systejti of reproduction. cell as The term cell is it is common now used traits have tranthat the for the metazoal well as for the microbe. the cells of an organism. macromolecules have to work in co-operation. One category. cells of a multicellular organism differ from the microbial cell which is an organism itself. In this way. macromolecules endowed with life a specific structure and a specific function. The and reproduction is the Both the cell of a multicellular organism and the microbe are the ultimate units of integration and reproduction. Reproduction. dependent. they many essential features that the scended the differences. cell. and Assimilation Life. The differentiated cells are organized into and the tissues into organs. corresponds to parts of an organism. as a controlled by the organism whole. is sometimes is listed among the discriminative features From a biochemical point of view.INTRODUCTION TO BIOLOGY and from the brain tissues cells. This can be accomplished only because of the presence and activity of specific templates that are parts of the living system.

prerequisite for assimilation. Y. M. (1911). A Symposiimi on the Cheinical Basis of Heredity. 269-3 10.Bioche?nistry. Reproduction includes assimilation. R. New The Cell . (1957). Englewood CliflFs. Antony van Leeuwenhoek a?7d His ''Little Animals. to synthesis. (1959). Protistenk. and the interplay of and functions. J. true to type and to genetic continuity.BIOLOGICAL ORDER The formula Metabolism is "the living being reproduces its its kind" expresses the fact that each organism produces a own specific macromolecules. York and London. What Is Life? The Physical Aspect of the Living Cell. Nature of Life. McElroy. (1948). A. is and reproduction. E. The University Press. Prentice-Hall. A. Szent-Gyorgyi. C. an organism can reproduce only if it is endowed with the power to assimilate.. W.. Stanier. New York. The Microbial World. (1944). and Adelberg. Physiology.. In the last analysis.. C. B. assimilation is certainly a discriminative feature of Assimilation lation. The Johns Hopkins Press. the notions of learned concerning life. organism. [6] . Morphology. Press. C. Sons & Danielsson. Schrodinger. In sense. Dobell. re- production. correlative to life. when something has been Anyhow." John Bale. (1932). eds. Doudoroff. It corresponds. growth. But the words reproduction and asshnilation will take their full significance only biological specificity. eds. E. (1957).. E. D. C. and Glass... London. The principles of protistology. J. Baltimore. and life is correlative to assimi- As a matter of fact. and Mirsky. Academic Dobell. and assimilation cannot be separated. Cambridge.. Inc. this life. 23. at the molecular the level of the organism. Academic Press. Arch. A. assimilation precludes specific structures level as well as at and includes metabolism. N. REFERENCES Brachet.

grows. This is the basis of evolution. etc. absorbs food. of one. Thus the growth and division of a bacterium is not the growth and division of its molecules. but a molecule cannot divide. same building blocks. nutrition. A molecule can be split into fragments. But we are not interested in the external appearance of things. metabolism. and divides. and can only be.Molecules might aggregate. which is two a microorganism. The specificity of an organism in the last analysis is. The statement [7] . This means that each individual always produces identical macromolethese species are constructed with the cules. What is reproduction at the molecular itself level? The But all living world is composed of thousands of species differing in their size. in the invisible essence of things. in We are not interested what we see but in what we do not see. . shape. some of which are inheritable. What is is it is biological specificity? How is the information stored which responsible for the biological specificity and diversity? How reproduced? Everybody knows that organisms undergo variations. A bv molecule is the smallest unit quantity of matter which can exist and retain all the properties of the original substance.THE PROBLEMS Organism and Molecules A bacterium. This process is metabolizes. but each fragment is necessarily different from the original structure. Neither can a molecule sfrow. the consequence of the specific structure of its macromolecules. bacteria are thus called reproduction' bv binary produced out fission. chemical composition.

Molecular freedom viral diseases are a catastrophe. in a sense fortunate. In harmony. that dangerously threatened. the constitutive. As a matter of fact. [8] . of its own As a result. about the way the organism as a whole decides what it has to do when faced with the genetic material. for otherwise life would be boring. Yet molecule sometimes decides to get rid of the shackles of co-ordination profit. In an ideal molecular society. all is sorts of molecular diseases are produced. When a molecule develops strong personality and refuses to submit herself to the common rule. Freedom is essential for the development and blooming of the human being. tion. How has the organism solved the problem of intermolecular communication? An organism rarely lives in a constant environment. is. are submitted to a dual action: they are controlled which is responsible for heredity and evolusame time they are subject to the action of the environment. of course. Each molecule Each molecule must be able to receive messages and must be disciplined enough to obey orders. its molecules. And one would like to know something about the interaction of hereditary and environmental factors. molecules have to work in know what the other molecules are doing. a what happens normally. When the environment varies.BIOLOGICAL ORDER that an organism reproduces true to type are is therefore in true only statistically. the organism has to cope with the variations. for the organism. Everyone knows This is that things are not always as they should be. dependent parts of the organan organism. molecular life. What the molecular basis of hereditary variation? An has to organism all is an integrated unit of structure and functions. and to work for diseases. the organism This is is each molecule works for the community. its experience. historical The structure of the organism includes past. But as part of an organized society the human being has to accept a certain number of restraints. I mean. Molecular a which only one aspect. by and at the a given situation. pose number of problems that we shall have to discuss. Heredity and is variation combined its the living system. ism. a This is true for molecules.

It was from the enzymes can be extracted only one chemical performs thus learned that a given enzyme to be catalyzed by reaction. and the word order has hardly been pronounced. The energy needed for the syntheses is provided bv^ the oxidation of food. and the energy obtained is stored as high-energy bonds. As everyone knows. is Biological order dual. In order to synthesize tryptophan. Syntheses take place. activity by themselves. A living being a dual system of order. We A have to know what this dual system is. each reaction requires energy. — COOH. Biological and it is especially a sequence in space a?2d time. synthesis of each building block is a stepwise process. Food is is seeded Chemical reactions are performed. Metabolism bacterium in. respectively. each enzyme. and finally each atom is exactly where it belongs. One atom is attached to another. sequence or succession in space or time. is taken in a proper nutrient medium. orders are is static and dy- namic. as a Order may also all that. for example sugar. carboxylated. a — CHo. for example in the form of adenosine triphosphoric one again mediated by a specific acid.THE PROBLEMS The Subject Our subject is biological order. Structural and functional the complementary aspects of the living being. The organism "burns" sugar. In a biosynthetic chain of reactions. Other enzymatic Some proteins have an help of a specific coenzyme. known concerning is the synthesis of or a intermediary metabolism. each reaction has a specific enzyme. [9] . As an The example. order is the fixed arrangement present be considered order is in the existing constitution of things. or aminated (attachment of. Most of enzymes work only with the the cell and purified. The oxidation of food and the storage of energy involve a series of chemical reactions. is Today. This the metabolism. molecule methylated. structural and functional. almost everything building blocks. a or an —NH2 in group). Enzymes are specific proteins. the biosynthesis of the amino acid tryptophan is depicted Figure 20.

what marvels there are in so small a creature." he wrote. "Dear God." The fact that life could not be something simple has been suspected for almost a century. In fact. the state of our knowledge. and size increases. The bacterial machine works. Claude Bernard produced an impressive statement. or perhaps the state of our ignorance. the bacterial cell. is life. they were cubes. a thousand packed into 1 milliliter. "by which one attempts to describe the protoplasm. Anyhow. cell the problem seems formidable and the situation hopeless.000 species of macromolecould be is one thousandth part of a millimeter. In his famed classical book. that is. And. itself divides. But for us. as the bacterium is devoid of brain.* corresponding to a volume of 10~i2 milliliter. not be concluded that knowledge and ignorance are synonymous. 1 /i A contains some 2.of them. machine of around 1 diameter. thus giving rise to two Simplicity or Complexity a well-known physical chemist interested in wrote this remarkable sentence. t If billions. manufacturing more of their far from simple. and finally the bacterium daughter cells. Les phhjomenes de la vie cormnuns aux animaux et aiix plantes {The Phenomena of Life Commoji to Afiimals a?id Plants). t a few thousand specific molecular species are at work. Claude Bernard certainly had no idea of how much more complicated protoplasm really is. synthesizes. however. And we would rather be inclined to say with Antony van Leeuwenhoek. is illusive. who in 1676 discovered the bacterial world. grows. lO^. the suffering human beings who try to A few years ago bacterial physiology penetrate the intimate nature of small." This was. [10] . In this specific kind. The nucleus divides. it has no problems. "The formula C18H9NO2. around 1875. Unity At * first sight.BIOLOGICAL ORDER As a result of the functioning of the enzymes. the protoplasm is much more complicated. the bacterium its synthesizes more enzymes. It should. "The structure of the bacterial cell is simple. despite being fx. and divides." This is true of course from the bacterial point of view.000 to 5.

Nucleic acids are made of four nucleic specificity is What will be learned later. the metabolism essentially the same in each Unity of coviposition: the main macromolecules of all living beings are composed of the same small molecules. nature has made use of a strictly limited number of building blocks. only necessary to For the time being. the problem of heredity. Uvity of pla?i: cules. in order to build the immense diversity of the living systems. each cell possesses a nucleus imbedded in protoplasm. know that the . gories of different organisms. one discovers unity. The problem of diversity of structures and functions. Yet.THE PROBLEMS Moreover. nature has produced an immense variety of catewhen the living world is considered at the cellular level. it is most important macromolecules are nucleic acids and proteins. and the prob- lem of diversification of species have been solved by the elegant use of a small number of building blocks organized into specific macromolecules. is Ufiity cell. of functiojT. For.

composed of twenty its An amino acid is.BIOLOGICAL ORDER bases. aminated acetic acid. for example.I \ C— C — C — COOH H N c H H NH2 V" . an aminated Glycine. In proteins./ " C^ I. as indicated is by name. thus forming nucleotide. thus forming long chain (Figure The amino acid. by phosphodiester bonds. analysis of proteins reveals that they are acids. A molecule of a phosphoric acid attached to each sugar. each bound to is a sugar to form a nucleoside. a The nucleotides are united 1). H H " \.

THE PROBLEMS molecule is for doing precisely endowed w ith a specific function. We may admire it. the living system did not perform its task. >/ [13] . The machine is built what it does. but we should If not lose our heads. We have simply to learn how it performs task. its it would not exist.

Schrodinger. in this fundamental book. tree is The a living reality. the chromosomal fiber. his well- known Esthetic. In its germ the determinants which was not already determinants exist as potentialities. the British philosopher Herbert Spencer. says the German philosopher. I have a suspicion that of the stu- somewhat unfamiliar with Hegel's Esthetic. Starting from small molecules — wrote Schrodinger in 1944 — it is possible to build large aggregates without the dull device of repetition. in his Principles of Biology. stated that in every organism the total of hereditary properties is determined by distinct physiological units which are formed by the assemblage of chemical units into immensely complex compounds. may be called an aperiodic [14] .THE HEREDITARY ORDER: GENETIC INFORMATION Birth of the ' Modern Concept Hegel published a small fraction In the year 1835. Whatever the case may be. even with the microscope. In 1864. and yet Nothing is in the germ one does in the tree not see anything. among a number of irrelevant remarks. every stone. in the germ. a crystal. The most essential part of a living being. the problem of dents here could be life is pertinently discussed. presenting the same solidity of structure as a crystal. plays an individual role not entirely equivalent to that of the others. Yet the most extraordinary biological intuition was that of a physicist. The atoms forming a molecule are united by forces of exactly the same nature as the numerous atoms which build a true solid. irrelevant for us today. as existing in the We can visualize the germ as extremely simple forces. In a complicated organic molecule. every group of atoms.

and not until 1953 that its it disclosed. The tions. how a molecule of hereditary material reproduced? Necessity of Variation The true. or ides. they were rediscovered inde- Hugo de Vries. is Each ide con- tains determinants. composed of biophores. which contains the chromosomes. according to They are relatively large Weismann. But Mendel's laws in remained ignored pendently by place on the material until. The chromosome. which means "colored bodies. and they stain heavily — hence their name. which all is composed of spherical bodies. in 1892. that to the unit of genetic material. Chromosomes have been known to cytologists for a long time. represent a different character. and each determinant is. Schrodinger adds that we believe the whole chromosomal fiber to be an aperiodic solid. And Weismann. that to multiply the specific biological order. would probably have disappeared long ago as the result of the drastic changes undergone [15] .THE HEREDITARY ORDER: GENETIC INFORMATION crystal. This was the beginning of formal genetics. and Tschermak. The biophore corresponds to the actual gene. But identified structure was was Each gene was assigned a specific was not until 1944 that the genetic chemically. is the cellular organelle which is responsible for the maintenance and transmission of the inheritable characters. Correns. understood that the inheritable characters were located in the chromosome. chromosome. Haeckel realized that the nucleus. and easy to stain. It is. the unit of life — has been defined as an independent system of integrated structures and funcis a highly complex system. able to reproduce its kind. If its kind is of course but only the primitive living being had life always given birth to an identical system. What is is the molecular basis of biological order and specificity? is How order stored in the is organism? If a molecule unable to divide. living being — the organism." As early as 1866. statement that the organism reproduces statistically true. The quantitative study of the transmission of hereditary characters in had been started 1865 by Gregor Mendel. 1900. the unit of biological order.

which a I would dislike to call final. which splits lactose into glucose and galactose. that is. quite obvious individuals belonging to one species were. Each time such situations it is found that the gene mutation. hereditary change. identical. or the hereditary change. are unable to oxidize amino acid by the normal route. a plant producing red flowers may modified offspring. some human mutants. When examined. Some human beings. responsible for the stepwise synthesis Human this beings normally oxidize the amino acid phenylalanine. phenyl- [16] . which is Mutations still take place nowadays. that if all It I suppose. It is admitted that the essential cause of evolutionary variation has sudden. later. as highly improbable systems. can only be the result of a stepwise process. found that one of the enzymes missing. Or a bac- terium synthesizing the amino acid tryptophan gives are analyzed. is acquired or is lost. evolution. place between the beginning. again. is. producing white flowers. wild type of another bacterium synthesizes the amino acid methionine. Living beings. the ability to synthesize a given enzyme. The biosynthesis is mediated by a series of enzymes. or mutants. the so-called a given bacterium is able to synthesize the enzyme ^-galactosidase. As a consequence. a given specific catalyst. there would be no such thing as genetics. corresponds to an alteration of an enzymatic system. Here the bacterium may produce mutants it is is unable to synthesize methionine. But wild type. Each enzyme corresponds to a specific gene. on the study of differences. rise to off- spring unable to synthesize tryptophan. or let us say an initial and a given. And the change is hereditary. As a result of a mutation. been the gene mutation. This the Beadle-Tatum law: one gene controls the synthesis of one enzyme. would be difficult to understand heredity. of it may The give rise to mutants devoid of this enzyme. Evolution is the sum of the alterations which is which have taken primitive state. The original. The principles of genetics are based differences.BIOLOGICAL ORDER by the primary environment. of transmissible give rise to For example. Let us give a few concrete examples. and would it is remain. In the absence of mutations. is transmitted to the offspring. So most fortunate that mutations do it exist. each responsible for a given step in the sequence of reactions. present state.

nature of the glucidic subunits are The polysaccharides are easy to identify. polysaccharideless bacterium. the Piieimwcoccus.Q. that to identify the nature of the entity controlling the production of an enzyme. unable to think. responsible for the the phenylpyruvic Long before the specific alteration responsible for a mutation had been chemically known. Material Identified Transformation. of these persons is is. The Genetic capsule. or which is a specific polysaccharide. They are idiots. This was accomplished by the study of bacteria. generally lower than 10. for a polysaccharide injected into an animal elicits the formation of a specific protein called an antibody (Figure saccharide combines specifically with the polyand not with polysaccharide II. the site of the biological alteration. Alany bacteria possess an outer layer. called ge7}es and described the of The problem was of course is. had been located on the chromosome. a Piieim703). and this information can be transferred to a bac- [17] . and long before the specific responsible enzyme had been identified. originally type II bacterium into a capsulated type III. The extract the of type III contains the "information" for the synthesis of type III polysaccharide. This however. originally of type in the for example. not important the in this analysis. Something present in extract of type III bacteria has trmisfoiined a capsuleless. The responsible as units chromosomal structures were variation. The agent of pneumonia. What is important is that enzyme present in normal human beings and normal pathway of phenylalanine is missing in idiots. produce its specific polysaccharide. is grown presence of extracts of type III bacteria. Sometimes. as already stated. The I. exhibits many varieties or types. It agglutinates bacteria of type III and not of type II. a mutation. The capsule of each of them is built of a specific polysaccharide in which the difl^erent. An anti-III-antibody III cocciis loses the ability to this capsuleless. to identify substrate of the hereditary variation. as a result of a sudden change.THE HEREDITARY ORDER: GENETIC INFORMATION pyruvic acid appears in the urine. When II. one recovers capsulated bacteria of type III.

BIOLOGICAL ORDER -^Antibodies agoinst type m ' ' .

endows it with endows it with a nucleic acid of given physiological potentiality. Desoxyribonucleic acid. Many bacteria have been transformed with DNA coming from a closely related form. multiplied and regularly transis But this specific nucleic acid not directly for the synthesis of the polysaccharide. a transforming principle can be extracted again. con- three types of molecules: (a) purine and pyrimidine bases. Avery. When a pure." This principle could not be metaphysical. The Finally. In all the cases of transformation studied so far. Thus the the specific is transforming principle. in a obtaining was Each fraction was tested for its transforming ability. the bacterium produces a specific a enzyme that can perform specific reaction. This is not a unique case but one particular example of a general phenomenon. it had to be an organic molecule. desoxyribose. MacLeod. or DNA. Once a gives rise bacterium has been "transformed" by a specific DNA. From the them. or DNA. [19] . They duplicate it in turn. and McCarthy succeeded in pure active substance that was identified as desoxyri- bonucleic acid. This information was called the "transforming principle. nucleic acid controlling synthesis of a specific substance. viruses can be separated The a from the protein components. that is. it to transformed offspring. Structure of the Genetic Material Desoxyribonucleic tains acid. in this case the synthesis of a specific poly- saccharide. the introduction of the desoxyribonucleic acid of the donor bacterium into the recipient organism the capacity to synthesize a given enzyme. specific viral nucleic acid penetrates specific viral proteins and specific viral infectious particles are produced. When it is present.THE HEREDITARY ORDER: GENETIC INFORMATION terium that was originally type II. and (c) phosphoric acid. naked. mitted from responsible cell to cell. bacterial extract containing the transforming principle fractionated. cell. The nucleic acid of viruses plays the same role in heredity as the nucleic acid of bacteria: both control the synthesis of specific proteins. 1944. (b) a pentose. The daughter bacteria have inherited the specific information. This conclusion has been con- firmed and extended by the study of viruses.

BIOLOGICAL ORDER

The two
idine
bases,

purine bases, adenine and guanine, and the

two pyrimFigure
4.
1

thymine and cytosine, are represented
3.

in

Each base

is

attached to a molecule of desoxyribose in position

carrying a phosphorus in position

This

is

a unit, a

desoxyribonu-

cleotide (Figure 1). In the nucleic acid, nucleotides are linked

by
1.)

phosphodiester bonds.

The

result

is

a

long chain. (See also Figure

adenine

thymine

guanine

cytosine
H

-H-K

'

H

\^~^cN
('sugars
C

\-/
C
//

\-H/

{ \

Vh
/

\
O'^UGAF

nh

Figure

4.

The Four Nucleic

Bases in the Desoxyribonucleic Acid.

is

Each pvrimidine base is bound to a purine base by hydrogen bonds. Adenine bound to thymine, guanine to cytosine.

Thanks

to

James Watson and Francis Crick, the structure of

DNA

has been disclosed.

DNA
is

is

a

double polynucleotide chain.

The two

by hydrogen bonds between the bases. bound to a purine base: adenine to thymine, guanine to cytosine. Thus in the double DNA helix, the ratio of adenine/thymine and of guanine/cytosine is equal to 1. The two
chains are held together

A

pvrimidine base

chains run in opposite directions.

They

are twined

around each

other (Figure 5).

[20]

THE HEREDITARY ORDER: GENETIC INFORMATION
This structure
is

now

firmly established.

The

evidence derives
(titration

from X-rav

studies as well as

from physical chemical data

curves, light scattering, viscosity, sedimentation, kinetics of break-

down by gamma
is

rays and enzymes).

Thus desoxyribonucleic acid is a double helix. Each helical chain made up of a chain of desoxyribose phosphate with one base
Ribonucleic
acid.

attached to each sugar.

DNA
is

is

the genetic material of
a

all

animals,
viruses

plants,

microorganisms, and of some viruses. But in

few

the genetic material

ribonucleic acid. Ribonucleic acid, or

RNA,
it is

has the same fundamental structure as
nucleotides.

DNA:

it is

is

a

long chain of

But whereas the sugar

in

DNA

desoxyribose,

ribose in

DNA
double

is,

RNA. Moreover, the pyrimidine base in RNA, replaced by uracil. Finally,

thymine present in whereas DNx\ is a

helix,

RNA
The
by

is

a single helix.

The two
information.

nucleic acids,

DNA

and

RNA,

both carry genetic
singularity

substrate of genetic information can only be a

feature shared
is

these

two

nucleic acids.

The common

obvioush" the linear arrangement of nucleic bases.

The chemical identification of the genetic material by Avery, MacLeod, and McCarthy has been the great discovery of modern biology. It was soon followed by another great discovery, that of the molecular structure of the desoxyribonucleic acid, the WatsonCrick double
virology,
helix. Both discoveries catalyzed the extraordinary development of genetics, cell physiology, cell biochemistry, and

which have now merged

into a

new

integrative discipline:

molecular biology.

Figure

5.

The Watson-Crick Model

of the

DNA

Double Helix.
are constituted

The two

chains run in opposite directions.

The backbones

by

desoxyribose molecules united by phosphodiester bonds. The nucleic bases of one chain are united to the bases of the other chain by hydrogen bonds.

[21]

BIOLOGICAL ORDER

The Molecular
Enzymes
species

Basis

of Enzymatic Specificity
are proteins. Proteins are
If,

of amino acids (Figure 6).
are organized

into

composed of twenty species two protein molecules, these twenty one and the same sequence, the two
in

Figure

6. The Molecule of Ribonuclease (from The Molecular Basis of Evohitio?i).

C. B. Anfinsen,

molecules are necessarily identical.
tein
is

The polypeptide

chain of a pro-

folded, but

it

will be admitted that folding

depends on the

sequence of amino acids and that two identical polypeptide chains

[22]

A given amino acid. and of course in the proportion of each of the twenty molecular species. except the terminal one. Fragment of a Polypeptide Chain. etc. at a specific peptide of the protein. Thus. is necessarily unique. The unit of function is the whole protein molecule.THE HEREDITARY ORDER: GENETIC INFORMATION same way.h CHg H HCH H CH2 H III CH2 CHg I I COO- CONH2 CH2 I \l + H3N^ I CH. differences in proteins in the sequence of amino acids which constitute the polypeptide chain. A long polypeptidic chain can be arbitrarily subdivided NH3 + OH ' CH2 C^ ' A 1 CH ' CH2 CHg I f-H 1^1 I c. When an enzyme acts. is necessarily associated to its left and to its right with another amino acid. the same peptides is may be repeated. being a specific molecule. Many molecules of the same amino acid may be present in a given protein. For the same groups of amino acids. the substrate attached at a specific site. the molecule. Their functional value depends on the sequence of all amino acids. In a protein. into peptides: di-. Each enzyme. tetra peptides. c K/ c CH2 H \c C II \ / c C II H \l ")c H \l c I c C ^C II — OH I N — OH I N —N OH I C II N I C II N I N II OH glycyl OH glutamyl OH III leucyl isoleucyl tyrosyl cysteinyl asparogine Figure 7. thus forming a tripeptide are necessarily folded in the can be due only to differences (Figure 7). no function can be assigned to any isolated amino acid or group of amino acids. But this specific peptide. This does not mean that in different species the same enzy- [23] . individual if isolated from the rest of would not be enzymatically active. tri-.

is antigenically (structurally) different in bacteria. If only one. synthesizes only a limited number of proteins. or group of closely related species. If just a were enough to take care of one amino acid. The enzyme /3-galactosidase. Now a protein is a specific sequence of amino acids. the The statea ment that each enzyme is unique applies only to the structure of given enzyme of a given species. in yeasts. a practically infinite array of specific proteins can be synthesized. is important is that proteins are synthesized on a nucleic acid plate. for example. and that is too low. Protein Synthesis: The Code DNA. with twenty different species of amino acids present in various numbers and proportions. One problem. with the is exception of some viruses. however. thus contained in has first seems probable that DNA to manufacture an RNA What tem- template and that proteins are synthesized on this template. A given organism. step by step. An organism is apparently unable to manufacture incomplete proteins. of the amino acids entering into the constitution of a protein is missing. and in animals. hydrolysis of the ^-galactosidic link. How does a sequence of four nucleic bases code a sequence of twenty amino acids? The problem of the code has been often discussed in recent years. If we assume that the process does not [24] . Things happen as if the synthesis were a sort of zipper-like process. It should be added that the synthesis of a protein is an all-or-none any one. starting at one extremity of the template and continuing process. The tem- plate therefore has to be specific. the number of permutations would be 64. the number of permutations would be 16. no protein synthesis is possible.BIOLOGICAL ORDER matic function cannot be performed by different proteins. We shall closely follow Francis Crick's excellent review. Let us pair of bases consider a sequence of nucleic bases: ACBDACDBCAD start at the C C B A C. has many solutions. It The highly hereditary information for the synthesis of proteins. Thus. If three bases were required and if any set of three bases could take care of one amino acid.

A A given amino acid can be coded only on one of "good triplets" being twenty. ACB.. the difficulties are the same. valid. BCA.THE HEREDITARY ORDER: GENETIC INFORMATION end. the total number This means that 44 triplets out of 64 do not make sense. the possibility^ that a to a is given sequence of amino acids. on one of the chains rectly built only. given triplet corresponds to only one amino acid. Moreover. If one is composed exclusively of "good" triplets. A given sequence of bases necessarily gives this hypothesis. ABA. ho\. and the end results will be determined by the point of departure. The good 8. Crick's Tentative Solution of the Coding Problem. is Any other does not others are bad triplets. Thus no overlapping rise possible. ABB. In single chain. according to the discussed code. etc. Crick. But the double DNA helix is from two complementary chains. No bining the adjacent bases of is good triplets can be obtained by comtwo good triplets. — which make sense — are represented in Figure All the A B C D Figure 8. the process can start in A or in C or in B. If there is a possible overlapping. Griffith. template a The protein may be synthesized di- on DNA is not excluded. Any make combination sense. and Orgel have worked out 1. the other chain would necessarily have a large fraction of "bad" triplets and would therefore not make sense.do the amino acids is know which template to read? If there no overlapping. a code possessing the following properties: triplet. there are 64 permutations. it would therefore be. . If proteins were synthesized on a DNA template. triplets 2.

even more refined data were oba bac- tained. has been learned that the nucleic acid is the substrate acid. they mate. however. Matthaei. Whether a sequence of three or four nucleotides is involved is not yet known. the coding problem considered open. Academic Press. ability of Two closely linked structures have a high probIt remaining joined. The spontaneous mutation rate in a bacterium or in * Since the Compton Lectures were delivered (March. 15881602]. pro"messenger. It is clear. By the use of mutagenic agents. is Anyhow. Moreover. work- ing with bacteriophage. 1961. DNA RNA [26] ." /. becomes attached onto a nonspecific cytoplasmic ribosome. New York. Sci." This messenger. template a. Thus: (a) the genetic information is contained (b) the code is a sequence of nucleic bases. that the whole code will soon be deciphered. Nirenberg and J. It has been learned that the structural gene. The code is now being deciphered: a synthetic poly-uridylic acid contains the information for the synthesis of a protein having the characteristics of poly-L-phenylalanine. our knowledge concerning the synthesis of proteins in general and the code in par- ticular has increased considerably. 1960). b. 47. 3. Proc. that the unit of variation was of the order of three to five nucleotides. to put in order). MoL Biol. of mutations. It is the system ribosome + messenger which assembles the amino acids into a given sequence: diataxy (etymologically. Acad. 318-356.* Mutation The nucleic it in the nucleic acid. detached from the chroduces a specific mosome. The longer the distance.6 to 2. is. A synthetic poly-cytydilic acid takes care of L-proline [M. the greater the probability of recombination. See Jacob and Alonod: "Genetic regulatory mechanisms in the synthesis of proteins.BIOLOGICAL ORDER Finally. the specific carrying the genetic information for the synthesis of a given protein. W. and Jacob and Wollman: Sexuality and the Genetics of Bacteria. thus producing a specific protein. When two same cytoplasm. It happens that the probability of recombinations is related to the distance between the two difl^erent chromosomes multiply in the factors. (1961). was found by Seymour Benzer. H. and recombinants are produced.8. How this is re- flected in the constitution of enzymes is not yet known. as entirely for the time being.'Nat. that of hereditary variation. The smallest unit of gene variation can be measured by recombination experiments. varies the ratio (adenine + thymine)/(guanine + cytosine) with the bacterial species from 0. (1961).

THE HEREDITARY ORDER: GENETIC INFORMATION relatively low: one ov^er one hundred thousand is (10~5) to one over one hundred million (10~8) per generation. Among these "mutagenic agents" are X rays.-. or naked viral nucleic acid are exposed to process nitrous acid. cytosine pairs with guanine.CH3 ^C_N > N^ J:c /O C H-N /C C /CH3 '. such as nitrogen mustard. This means that the deamination of one pair. an adenine/thymine of deamination._/ \ \ / ^N r q/ C \_N —H ( hypoxanthine deam. A4any agents are known to increase the probability of mutation. The is kinetics of the a one-hit process. base enough to produce one mutation. or animal viruses. which is thus converted into hypoxanthine. or bacteriophage. bacteria.ne) thymine CH3 H N deamination \ / H \ hypoxanthine H H cytosine guanine H cytosine \ C— N ^N^. noted oden. The original base pair adenine /thymine has a result been replaced bv the base pair guanine/cytosine as of the action of the nitrous acid. At the second rephcation. Nitrous is. >— . mutants are produced. for example. and teriophage nitrous acid. deaminates nucleic bases. ultraviolet rays. and various chemicals. Let us consider. H Odenme thymine N ' >-< < }-< "-\_/"^" >-% /-" \ /"' \/^-\ . organic peroxides. adenine 9). as will be seen later. when is acting on nucleic acid. that takes off their NHo group. When. or plant viruses. is As a result transformed into hypoxanthine (Figure The replication of the DNA double helix. [27] .0 H —N / C / C N— H Z ^N >—< N \—N . The Mutagenic Action of Nitrous Acid (from Robert Lavalle). reveals that induction of mutations acid. At the first replication. for example.Cy <.CH ^c )^-" \ C/ /" "^ \-H after one replication and tautomerization after a second replication Figure 9. Nitrous acid deaminates adenine. hypoxanthine binds cytosine.

" The "mutated" protein? protein. to a given wild-type gene and to this gene once has undergone mutation? Hemoglobin. nothing more. of a complementary strand. the original base pair. By But the differences have been traced back to one amino acid. Amino Acids Sequence Hemoglobins. the hemoglobin can be cut into small peptides. which in turn will bind guanine. In a hereditary blood disease of man. the Table I. the protein of the red blood of oxygen. one of these peptides is different in the normal and mutated proteins. Thus. What the result of a gene mutation in the What a is the difference between the proteins corresponding. the pair guanine/ is one of the daughter cytosine. hence the sickle name of different the disease: sickle-cell anemia. replaced by in fact a so-called "point mutation. will bind cytosine. hypoxanthine. is in helices.BIOLOGICAL ORDER essentially the formation. As discovered by Vernon Ingram. mild digestion. in Normal and Abnormal Peptide fragments of normal hemoglobin (A). The normal human red cell is a disc: it looks round when it lies on its flat surface. associated with the so-called "hemoglobin C disease. The in a hemoglobin is from the normal hemoglobin number of properties (Table I). has been the subject cells responsible for the transfer of extensive studies. The new base. and hemoglobin (C). We is have learned that nucleic acid con- trols protein synthesis. In the sickle hemoglobin S. A gene mutation." Histidine . by each of the strands. the red cells are reduced in number and exhibit the form of a sickle. as a consequence of a deis amination by nitrous acid. it respectively. adenine/thymine. sickle-cell hemoglobin (S).

THE HEREDITARY ORDER: GENETIC INFORMATION
glutamic acid
acid
is

is

replaced by valine. In hemoglobin C, glutamic
lysine.

replaced

by

Thus we have

learned that nucleic acid

is

the substrate of heredi-

tary properties, and that the genes control the synthesis of specific
proteins, or of specific enzymes.

We
is

know

that the sequence of

nucleic bases in the genetic material

responsible for the sequence

of amino acids in the proteins.
base pair

We know that the

replacement of one

by another may be
difference

responsible for a mutation.

We

also

between the normal and the mutated protein may be due to the replacement of one amino acid by another. As a consequence of the replacement of one base pair by another in the gene, one amino acid is replaced by another in the protein. The sequence is changed, and as a consequence the folding of the polypeptide chain might be changed too. The nucleic acid,
that the

know

the genetic material,

is is

the substrate of the structural, hereditary
the agent through

change.

The
is

protein

which the hereditary

change

expressed.

Reproduction of the Genetic Material

DNA. The
that
is,

genetic material, or the genetic information,

is,

by
cell,

definition, transmitted

from the mother

cell to the

daughter

multiplied.

How

does one molecule of nucleic acid produce

two
of

identical molecules?

How

has nucleic acid solved the problem

acid is a double and thymine on the one hand and guanine and cytosine on the other hand are united face to face by hydrogen bonds. The double helix is thus composed of two complementary strands. Watson and Crick in 1953 put forward the hypothesis
helix in M'hich adenine

molecular reproduction?

Desoxyribonucleic

that each strand acts as a template for the organization of a

comple-

mentary one.

As

the

tion of the
in fact,

two strands of the duplex are complementary, the separatwo strands cannot be described as a binary fission; it is, an inequational cleavage, the separation of two different

But as these parts are complementary, each one, when producing its complement, reproduces the original molecule. The unit,
parts.

[29]

BIOLOGICAL ORDER
which is the molecule, is made up of two firmly bound complementary parts. The molecule of DNA cannot grow and does not grow. But by making use of complementarity it has solved the problem of molecular reproduction. By the template mechanism, a single strand of nucleic acid can produce a molecule of protein. It can also produce a complementary nucleic strand, which in turn can reproduce the original one. But only a molecule made of two complementary structures can reproduce directly its own structure. This is all right on paper, but how are things in the living system? An enzyme has been isolated from a bacterium that polymerizes desoxyribonucleotides. If these building blocks of DNA are added in a test tube together with the enzyme, is synthesized. The necessary condition, however, is the presence of a DNA primer.

DNA

The
In

original

DNA,
as

the primer,

may

be multiplied

i?i

vitro

bv

a

factor of 10.

DNA,

already mentioned, adenine

is

bound

to thymine,

guanine to cytosine.
guanine/cytosine, (adenine
is

The
equal

ratio

adenine/thymine,
1.

like

the
the

ratio
ratio

to

But,

in

bacteria,

+

thymine) /(guanine

+

cytosine) varies from 0.6 to 2.8

according to the species.

What
primers?
First:

happens

when

these various types of

DNA

are taken as

Second:
missing.

The ratio adenine/thymine and guanine/cytosine is always The synthesis does not take place if one of the bases

1.

is

Third: Each base can be replaced by an analogue, provided its hydrogen-bonding capacities are in agreement with the Watson-

Crick model.
Fourth:

The

ratio

(A

-f-

T)/(G + C)

in the synthesized

DNA

is

the same as in the primer. If the "nonnatural" synthetic primer does

not contain guanine or cytosine, these bases are absent in the newly

formed
Fifth:

DNA, despite being present in the test tube. When the DNA primer is heated, the two strands
is

separate,

and the rate of synthesis
Everything happens

markedly increased.
if

as

one of the strands would act

as

a

[30]

THE HEREDITARY ORDER: GENETIC INFORMATION
template. Experimental data obtained so far are in full agreement

with the Watson-Crick hypothesis.

more? Three mechanisms could be envisaged for the replication of the DNA. These are called conservative, semiconservative, and dispersive (Figure 10).
learn something

Can we

1 a

U

form another pair of hydrogen bonds with a third base. an enzyme has been discovered which binds ribonucleotides and organizes them in long chains. Its DNA was labeled with radioactive phosphorus. The formation of a complementary strand by the template mechanism would end in two chains with qualitatively different genetic information. an experiment consists of labeling the is needed. As the two chains of the double DNA helix are complementary. and is more generally not yet solved. the synthesis of proteins. The same type of problem is posed by RNA. It DNA of the parent organism and following the distribution of the label in the offspring.BIOLOGICAL ORDER In order to answer our question. Reproduction of RNA. A number of experiments perform. One could of course be active. 3. Only one of the strands of the DNA duplex produces an RNA the chain. the C of the sugar and the N of the base is to be always in the same 2. Bacteriophage was selected as the material. But it could well be that [32] . The conclusion was that DNA reproduces by the semiconservative mechanism. position. according to Stent. Each of the two strands of the DNA duplex forms an RNA chain. specific sequence of the RNA bases Each of the bases of the base pair of the duplex could. in agreement with the original Watson-Crick hypotheses. a given pair of the duplex bases would combine with only one base of the future RNA. The single-stranded RNA The is formed within the deep groove of the is DNA double helix. the two RNA chains would be different. RNA is single-stranded. Three mecha- nisms have been considered. Production of an RNA template froiM DNA. If the bond between pairs of the governed by the base DNA duplex. The first experiment by Cyrus Levinthal in 1956. 1. which is the genetic material of a number of animal viruses and of plant viruses. The problem of the mechanism by which DNA produces the RNA template for an RNA molecule.ed since on various materials seem to agree with this of this kind was done conclusion. and the concentration of hot phosphorus was measured in the daughter bacteriophage. the other being inactive. that is. Here again. the other inactive.

. Anfinsen." The key problem now REFERENCES The Molecular Basis of Evolution. B. The Macmillan Com- New York. Cambridge. (1951). forget the question of the replication of the ribonucleic acid and of viruses containing ribonucleic acid.THE HEREDITARY ORDER: GENETIC INFORMATION RNA is A — possesses the right sequence of triplets coding of amino acids and the synthesis of proteins. or that the single-stranded produces a double-stranded DNA. E. The mechanism would be the reonly one — let us say suitable for the RNA RNA verse of the synthesis of RNA from us." wrote Max Delbriick in 1949. New Dunn. Genetics in the 20th Century. (1947). desoxyribonucleic acid. one may perhaps suspect that the key problem of biology. how living matter manages is and perpetrate experience. Dyna77nc Aspects of Bioche772istry. physicist's point of view. "that the essence of life is the accumulation of experience through the generations. York. Anyhow. Conclusions The conclusion has been reached that each organism contains a structure. the problem not solved — not yet solved. Baldwin. C. then. John Wiley & Sons. A change "If it in the base sequence results change in the nucleic acid specificity. L. This structure contains the information for the pro- duction of specific templates and of specific proteins. [33] . pany. C. is from the to record solved. be true. which is the very base of heredi- tary order. DNA. The University Press. The specificity of nucleic acid is is the sequence of the nucleic bases. The other — let us say B — would be able to produce only a complementary strand A. Let for the time being. its reproduction is the production by each of the strands of a complementary one. and the sequence of the in a the specificity of proteins amino acids. ed. Because is DNA composed of two complementary strands. It could also be that a temporary double helix is formed. (1959).

Baltimore. eds. John Wiley & Sons.BIOLOGICAL ORDER Fruton. (1955). D. S. (1959). York. 17-26. Rich. [34] . The Johns Hopkins Press. Wagner. McElroy. John Wiley & Levinthal.. (1956). 42. Genetics and Metabolism. K. Nat. A. C. New York. and Simmonds. The mechanism of DNA repHcation and genetic recom394-404. Brookhaven Symposia in Biology. bination in phage.. General Biochejnistry. R. Proc. Sons.. Set. Acad.. S. and Mitchell. (1957). 12. Polynucleotide interactions and the nucleic acids. B. J. H.. P. New W. A Sy7t7posium on the Chemical Basis of Heredity. and Glass. (1953).

A bacterium contains about 2.000 genes and 2. the building blocks are synthesized step — the nucleic bases and the amino acids specific proteins — by step. are interdependent subunits. the double mother divides into two daughter cells. This DNA is reproduction. exist If the factory manufactures cars. which. that is. a given balance must etc. the building blocks are assembled in an orderly way and specific nucleic acids. In the first phase. or RNA. or something. doors. cars. wheels. When the genetic material has duplicated. they are [35] . acids Each enzyme is a specific sequence of amino which performs only one chemical reaction. has to co-ordinate and to orient the activity of the individual units.THE FUNCTIONAL ORDER The Problem Life has been equated with a complex hereditary order invoh'ing a specific sequence of nucleic bases. For the first process. are parts of a whole. Growth and multiplication depend on the completion of two sets of processes (Figure 1 1 ). The second process involves specific templates. only enzymes are required. This is growth. that depends on extrinsic factors. as parts of a whole. Thus more enzymes and more nucleic acids are produced. The units is.000 enzymes. contains the information for the synthesis of enzymes. the genetic material. The fiber of nucleic acid. between the number of motors. some- body. In the second into phase. the workers. Aioreover. brakes. In a factory involving the activity of a few thousand workers. the rate of production has to be adjusted to the con- sumption of it of the factory.

the essential metabolites (amino acids and nucleic bases). It has to synthesize all the in the right proportions in excess and in the right amounts. are synthesized. And waste decreases the chances aminoacids metabolism reproduction Figure 11. and Reproduction. The duced cell is a factory. The statement that the organism reproduces true type is [36] . A substance proin means waste. Metabolism. An imbalance may lead to cellular disease and even to live in a death. Moreover. Nucleic bases are organized into nucleic acids. especially between nucleic acids and proteins.BIOLOGICAL ORDER interdependent units. Food is metabolized. organisms do not constant environment. the Struggle for life. In nature. This is true especially for bacteria. The small building blocks. they encounter a wide variety of conditions. these poor beasts to are submitted to terrifying ill-treatment. different types of devices conbuilding blocks trolling order are interacting at various levels. The duplication of the desoxyribonucleic acid (DNA) is is the very basis of reproduction. prerequisite for protein The manufacture synthesis. A balance also has to be maintained between the various macromolecules. Amino acids will of ribonucleic acid the be patternized into specific protein on the ribonucleic acid templates. Growth. In a factory. And in the laboratory.

the bacteria are unable to grow in a medium containing carbon and energy source. galactosidase is synthesized only when lactose or a few other ^S-galactosides are present. considered: inhibition of As an introduction. The potentialities of the organism are under the control of the subject to the action of the environment. the integrated system of specific interdependent macromolecules. If lactose is supplied. The enzyme that controls its metabolism splits the /?-galactosidic bond and is therefore called /3-galactosidase. The induced synthesis of the enzyme lactose as the sole [37] . quantitatively as well as qualitatively. As a result of its is activity. When no lactose is present. two models will be one of induction of enzyme synthesis. but the functioning of the genetic material and of the whole organism is The organism as to has to cope with the environmental changes as well its ensure the harmonious interplay of in the enzymes. tose. Bacteria grown in a medium containing Bacteria in a synthetic grown glycerol are completely devoid of enzyme. Lactose induces the synthesis of galactosidase. the synthesis of the galactosidase starts after a lag of a few minutes. We have to dis- is informed of cytoplasm and in the outer world. however. Thus. genetic material. This a prerequisite for the utilization medium where the only organic compound is lactose contain the enzyme galactosidase — about 6.THE FUNCTIONAL ORDER Statistically correct. Lactose is a which glucose and galactose are united by a ^-galactosidic link.000 molecules per bacterium. How are the orders transmitted from one macromolecule to the other? What is the basis of molecular interaction and balance? The cell. In the absence of galactosidase. disaccharide Models of Enzyme Synthesis of the bacterium Escherichia coli are Some in strains able to utilize lactose as a carbon or energy source. and within an hour or so each bacterium has manufactured its full load of enzyme. lactose is hydrolyzed into glucose and galacof lactose. Two Induction. the enzyme is useless. the other of enzyme synthesis. will be close the mechanism by which the genetic material what happens studied as a functional unit. The enzymatic equipment reproduced true to type of a bacterium is is the far from being constant. But what It varies is genetic material.

Of course they have medium. building blocks. that is. When growing in a synthetic methionine synthase. What happens when the growth in the absence of methionine. synthetic The bacterium Escherichia coli is able to grow in medium devoid of any amino acid. the bacteria contain to. they are unable to synthesize methionine. [38] . or essential me- among them. to © cysteic C C II SO. Such a bacterium all its therefore able to synthesize tabolites. why this It is process has been. C ^ ^ _ COOH COOH COOH C NHj C — NHj COOH C I S — CH3 c I C I — NH. for a long time. and. COOH homocystine methionine METHIONINE SYNTHASE Figure 12. called enzymatic adaptation. And they are unable to synthesize grow. The enzyme methionine synthase is actually present (Figure 12). a is Repression. proteins. The Synthesis of the Amino Acid Methionine. Let us now consider the reverse process. now described as an induced biosynthesis of enzyme. the amino acid methionine. is The last step in the synthesis of methionine catalyzed by an enzyme called methionine synthase. For in the absence of methionine synthase. NHg acid—H* cysteine " C "t" ^ homosenne C SH NH2 C S C— OH .BIOLOGICAL ORDER increases the fitness It is between the bacterium and its environment.

sponsible as a result is of this excess.THE FUNCTIONAL ORDER medium is is utilized. The cytoplasm is limited by a thin membrane. we observe an adjustment of the enzymatic The cell tends toward a physiological balance. the bacteria are assayed enzyme has disappeared. But. A balanced state to an equal production is reached which corresponds and consumption of methionine. is surrounded by a cell wall whose function is mainly counteract the internal The bacterium to osmotic pressure. The development of our knowledge concerning the functional order has been possible. after for methionine synthase: the few hours. is Suppose now that the bacterium it methionine. Let us consider next the problem of the interactions of genes and of enzymes. a decreased rate of is. Whether we deal utilization of the carbon and energy source. the inhibition of called repression. This excess can be due either to an excess of enzyme or to utilization of methionine. And. bacterium and its environment. The result is the same. namely the amino acid methionine. thanks to the existence of nonhereditary. Our problem is to learn what the responsible mechanism is. system. a is supplemented with methionine? A labeled methionine methionine with a hot sulfur atom. Among the remarkable properties of [39] . Repression of enzyme synthesis thus increases the the fitness between with the lactose. The labeled methia onine taken up. and that is growing in the absence of produces an excess of methionine. The Genetic Control of Bacterial Metabolism Metabolism of lactose. the synthesis of the re- enzyme repressed. Escherichia coli will be taken as a tool. namely the sugar or with the synthesis of an essential building block. such as the induced synthesis of enzymes or the repression of en- zyme synthesis. The development of genetics and the chemical identification of the genetic material have been possible because of the existence of hereditary variation called mutations. not transmissible variation. that an excess of methionine. This means that in the presence of extrinsic methionine. This phenomenon. and as a consequence the synthesis of methionine decreases. the synthesis of the enzyme is blocked. enzyme synthesis.

" [40] . The two structural genes y and z control the synthesis of the /3-galactoside permease and the ^-galactosidase. bacterium. another for the sugar glucose. Lactose the by the model bacterium possesses Both enzymes are synthesized is an inducer. etc. o D — I ' permease cr •^gal I • I 1+ z+ y+ ^ > O o v^ • O O • O ^ /3. Our both permease and /?-galactosidase.galactosidase Figure 13. as in logic. the i /3-galactosidase splits lactose into glucose and galactose. another for the amino acid methionine. enzyme called permease.BIOLOGICAL ORDER the bacterium is selective permeability. Its is controlled by a specific per- role is just to take the molecules of lactose in the environ- ment and to transfer is them it is across the membrane. Accordingly. split Once lactose has been pumped into the cytoplasm. a Schematic representation of respectively. into glucose and galactose original enzyme ^-galactosidase. The permease /3-galactosides is responsible for the specific transfer of lactose and other the across membrane. only when lactose is present. The Utilization of Lactose. "based upon or using induction. A bacterium is not permeinside able to everything. lactose -^glucose + galactose —n . (Figure 13). The gene controls inductivity versus constitutivity. enzymes are said to be inductive. Inductive means here. the bacterium of And it was discovered that the penetration many small molecules is mediated by a There is given specific one specific permease for the amino acid tryptophan. a reaction in which an acetylation perhaps involved. the metabolism of lactose mease. Thus. another one for lactose.

to rise to hereditary vari- mutants belonging to various categories: bacteria able to synthesize /5-galactosidase (a) The may give rise to variants unable to perform the synthesis. (f o . This original model can give ants. the mutation is z+ ±^ z~.THE FUNCTIONAL ORDER Variations. z+ being the symbol for the character carrying the information for the galactosidase.

Related Interests

^ = inductive I i+ I z+ y+ K I i i+ O = permease • = y5-galactosidase /5 o 1 O • 6 • • 6 • =constitutive . where i+ stands for inductivity. the mutation + i~. i+ in the absence of an inducer. being the symbol for the charis acter controlling inductivity versus constitutivity.BIOLOGICAL ORDER (c) The inductive bacteria may give rise to mutants that pro- duce /?-galactosidase and permease They i are called constitutive. i~ for constitutivity (Figure 15).

This gives a measure of the relative dis- tance of the genes. pair The normal gene ment of one base A point mutation. z+ . template. and i. b' I Figure 16. as a unit. mechanism. for example in a (a+ > a~) or in b (b+ > b~). As already Are seen. controlling the utilization of lactose. or corresponding of zipper-like 16). one finds: that the characters of the male can be transferred to the female.THE FUNCTIONAL ORDER The gene jugate. possesses the structure a+ b+. is the friction forces separate the partners. namely. 2. the determination of the frequency of exchange of one piece of a chromosome for another. Gene Mutation. When 1. and possible types of combination are found. as a whole. If this done at intervals. y+ . can mutate independently. [43] . a— b+ 1 mutation A „ mutation 8 . Let us supposed that a gene. When males are mixed with females. that the three "genes. the process lasting around one hour. can take place on various gene." z. and is conjugation interrupted. one finds that each gene of the male enters the female at a given fixed time. and that the various genes always enter in the same order. . This order can be verified by the classical tests of genetics. a replacesectors of the by another. these types of experiments are performed. these genes parts of the same functional unit. produces a protein. This allows us to locate the place of each gene on the chromosome. by a sort now consider the gene z (Figure The a+ b+ 1 normal gene . Our bacterium exhibits sexual processes. i+. or do thev reprethat the units sent different functional units? are different? How can we know its What justifies It is the use of the term "unit"? Altered enzymes. the all different genes. If the couples are put in a Waring Blendor. are closely associated or linked. they conof the male is The chromosome injected slowly into the female. sexual digression. y.

but instead a new. altered protein. to enzyme. Now the mutants a~ and b~ are different. the normal structure. The Activity of Normal and Mutated Genes. that is. that is. is present. It may have a slight enzymatic activity. that a fraction is. is present. one to the a+b+ corresponds for the the information synthesis of the Table II. In [44] . When the mutated gene z~. despite the presence of all the information. the enzyme is synthesized (Figure 17). The mutated gene z~ does not produce the original enzyme. Let us then mix the various z~ mutants. they can saturate (20 to 60 per 100) of the antibody.BIOLOGICAL ORDER structure "normal" gene z+. replaced by Some only fold. All this shows that the mutations affecting the gene z+ are mani- The proteins produced /3-galactosidase by the various z~ mutants differ from the enzyme and differ also in each of the mutants. As a result. others give incomplete cross-reactions. and any nucleic base. ture /?-galactosidase. Such a heterozygote is unable to manufac. can be replaced by another (Table II). The normal gene z+ produces the enzyme. Some z~ mutants give a complete cross-reaction. they are able to displace entirely the enzyme from the enzyme-antibody complex. whether a~ or b~. and in the protein one amino another. The male carries a gene a+b~ and the female a gene a~b + The zygote has two different sets of genes and is therefore called heterozygote for gene z. or be totally devoid of activity. An important fraction of the negative mutants z~ nevertheless contains a protein able to cross-react with the anti-/3-galactosidase antibody and to displace ^-galactosidase when combined with the antibody. whether in position a or b. no enzyme is synthesized. — normal enzyme (less "Normal" gene Mutation Mutation II I > active protein > active protein — modified enzyme — no enzymatic potent) > inactive protein activity sequence of bases acid is is replaced by another. So when the gene z+(a+b+). A gene is a sequence of nucleic bases.

THE FUNCTIONAL ORDER < d .

The male injects its chromosome female. Why that an inducer sometimes needed for the to gene to express tutive itself. When the bacterium necessary. it is One to takes a z+i+ female that possesses the right information for the synthesis of the enzyme. Another hypothesis has to be considered: the gene i+ manufactures a repressor.BIOLOGICAL ORDER Induction and Repression Generalized induction hypothesis. is i+ (inductive). is is is. then the zygote should synthesize the enzyme in the absence of an extrinsic inducer. and the female is z~i~ instead of the male. the enzyme synthesized whether or not an extrinsic inducer is present. the gene i~ being unable to Let us then admit that the gene i+ does synthesize a repressor. The repressor. the heterozygote z+i+/z~i~ starts producing /3-galactosidase at full speed. The reverse experiment is then performed. One takes a it z^i" male (Figure It cannot synthesize enzyme it because does not possess the right information. but. The into the male is z+i+ instead of the female. to inductivity. that is. if the function of the gene i~ is to manufacture an inducer. In the absence of an inducer. And the constitutive female cannot manufacture the enzyme because it is z~ . A [46] . an extrinsic inducer is. do so. unable males manufacture enzyme in the absence of an inducer. by definition. But possesses the constitutive gene i~. It tested. the female cytoplasm corresponds to the gene i+. could produce enzyme in the absence of an inducer if it were z+. manufactures an inducer This hypothesis can be easily 18). The gene z+ contains all of the information for the synthesis of a y8-galactosidase. Now and females are mixed. Male and as the female are again mixed in the absence of an inducer. Things are more complicated than was thought. make use of its potentiality is by manuit facturing the enzyme? The simplest hypothesis that the consti- bacterium does not need an extrinsic inducer because itself. When the bacterium is i"~ (constitutive). If the theory is right. It does not. In the zygote type I. the male cannot facture the manu- enzyme because it is inductive. But the bacterium that possesses the gene z+ does not necessarily manufacture the enzyme. being i+ (inductive). As soon z+i+ gene from the male has penetrated in the z"~i~ female.

THE FUNCTIONAL ORDER X X X X X ^ X X 2+ X X 5^ X X X X ^X X "" X ^ X X X > X X v- X I I cr zygote I .

should be responsible for the manufacture of a repressor. which. in is the absence of an inducer. This If. and the enzyme synthesized. is No repressor is present. But the male has introduced at z+ introduced the same time the phase I • X :/?-galactosidase = repressor • X • HOURS Figure 19.BIOLOGICAL ORDER repressor is is present. This the first phase. The by heterozygote of type (i~). according to the hypothesis. the synthesis of the enzyme gene If. is the second phase. however. an inducer is added. the female cytoplasm i~. being constitutive devoid of repressor. Something seems to be wrong. as soon as the gene z+ enters the female. Synthesis of /3-Galactosidase II by a Heterozygote. then found that the rate of enzyme synthesis decreases and reaches a zero value (Figure 19). This corresponds to constitutivity. thanks to the gene by the male. But the male has introduced also the inductive gene i+: is a repressor synthesized. the synthesis of galactosidase is followed long enough. it is [48] . continues. i+. and the synthesis of galactosidase stopped. and ^-galactosidase synthesized. In the zygote type II. shown in the preceding figure has been formed the transfer of the male genes i+ z+ into the i~ z~ female. A repressor has really been formed. however. is The is female.

And is it is quite natural that the synthesis of the contingent enzymes acids. genetic analysis has thus revealed a number of important The genes z. Thus. determines the constitutive as opposed to the inductive type of synthesis of both enzymes. lactose is a carbon source and a source of energy. The Ways of Repression Repressor. z and y. the bacterium stops synthesizing methionine synthase. One. It is called a regulating gejie.THE FUNCTIONAL ORDER If an inducer is tion of enzyme starts added after the synthesis has stopped. the control OF enzyme syntheses. Let us consider the enzymes responsible for the synthesis of the essential building blocks and especially of amino Amino acids. the hazards of to its The food daily life. ^-galactosides. The repressor has not yet been identified. is the biosynthesis of amino acid arginine. It is part of the food. They are called structural genes. But a number of data are in ^ favor of the hypothesis that it is a ribonucleic acid. It has already is methionine been stated that if the amino acid provided. Things happen as if the gene i+ controlling the inductive state corresponded to the i~ to the loss of the synthesis of a repressor. corepressor. y. For the bacterium. the small sector of the bacterial chromosome is that controls the synthesis responsible for the metabolism of lactose. This general. such as glycerol or pyruvic acid. and inducer. The bacterium may be fed with other organic compounds. of galactosidase and permease. the enzyme responsible for the last step in the biosyn- thesis of methionine. and a mechanism the known to be quite few cases have been studied extensively. Two carry the information for the synthesis. of the enzymes generally of more and of them. respectively. The product of enzymatic activity stops the is synthesis of the enzyme. aporepressor. i. that a bacterium depends on The bacterium obviously cannot afford find may manufacture a large number of useless enzymes. regulated. the mutation i+ ability to form a repressor. composed of three units. The facts. and i are independent units of function and therefore are different genes. hence to a constitutive situation. The other. the producanew. for In a example. [49] .

histidine and tryp- tophan. If supplied. The Biosynthesis of the Amino Acid Tryptophan. ® ® anthranilic acid ® (D tryptophane shikimic acid TRYPTOPHANE SYNTHETASE Figure 20. if an excess of arginine is produced.C. NH. Thus. such as nucleic bases. specific.BIOLOGICAL ORDER medium devoid arginine is of arginine. the synthesis of the enzymes responsible for the synthesis of tryptophan is not altered in the derepressed strain. In a synthetic synthesized by a medium devoid of tryptophan. repressed. and this excess tryptophan is excreted. Only the case of tryptophan will be discussed here. The produce much more tryptophan than needed. it is possible to isolate mu- tants that are called derepressed. Let blocked by a repressor. the bacterium synthesizes arginine. whereas it is inhibited the tryptophan enzymes bacteria [50] . If tryptophan is is is added. This repressor on the tryptophan enzymes and not on the synthesis of any other system. serine HO— C— C— COOH H I NHg COOH A. is and less arginine will be produced. the synthesis of is not blocked any more by tryptophan. In these mutants. HO- COOH /^ ® n indole C — C I COOH Y OH °" <d) NHg—••B. the synthesis of the enzymes involved in its synthesis is specifically repressed. the enzymes cease to be produced. If tryptophan is added to the medium. The same mechanism operative in a number of biosynthetic pathways. that is. the synthesis of the enzymes us admit that the synthesis is is stopped. as shown in Table III. The end product of the chain of biosynthesis blocks the synthesis of all the enzymes involved in the chain of arginine biosynthesis. it is known to act only From the original repressed bacterium.D. tryptophan system of enzymes (Figure 20).

c o D- u 4-> to (U c W c o c « Oh N c O I-H Cl- H c 'M o .

unable to manufacture the repressor. The without effect on the enzyme level in the derepressed strain.BIOLOGICAL ORDER Let us admit that the original wild type which is able to manufacture the repressor. is A obviously produced by the R+ gene. allowed to conjugate. Table IV. which dominant over the R~ gene (Table IV). Enzyme Synthesis as Affected by Tryptophan in Repressed and a Derepressed Bacterium and in the Heterozygote. the synthesis of the tryptophan repressor is enzyme is blocked. and that the derepressed bacterium carries a mutated gene R~ try. In this zygote is thus formed. The hetero- zygote behaves "repressed" is like the repressed strain: the character dominant. possesses the gene R+try. a The times derepressed bacteria contain three to seven the more enzymes than is repressed one. A and R~try genes. Without tryptophan . carrying both R+try heterozygote. R+ and R" bacteria are in the normal strain. whereas it blocks addition of tryptophan enzyme synthesis in the repressed one.

When growing in a medium containing glucose and lactose. but it expense of the reserves that the bac- A corepressor perhaps continues to be pro- has less affinity for the aporepressor than the product of glucose metabolism. The theory is that a product of the metabolism of glucose acts as a corepressor \'hich combines with an aporepressor. But is an excess of /^-galactosidase as a result synthesized. Lactose competes for the aporepressor with this hypothetical corepressor of low affinity and thus prevents is the formation of the repressor. But at the when it is used up. no /^-galactosidase start. consequently. The synthesis of the enzyme respon- for the metabolism of lactose will be repressed. In view of the relatively rapid adjustment of enzyme synthesis to the need.THE FUNCTIONAL ORDER Whether the tryptophan has been manufactured by the bacterium or pumped from the medium into the bacterium. is produced by the z+ bacteria. is glucose is used entirely. Things happen as if in the presence of glucose a repressor has been produced. A corepressor will be formed and. When it has disap- peared as a result of the cellular metabolism. an excess of glucose produced sible of its activity. a corepressor not produced or is is not available. duced. If The up if latter are used up in the metabolic processes. How does the aporepressor-repressor hypothesis apply to the regulation of enzymes involved in the metabolism of the carbon and energy source? When lactose is present as the sole carbon and energy source. the bacterium "needs" galactosidase in order to grow. it.8-galactosidase synthesized and functions: lactose is hydrolyzed into glucose and galactose. provided it possesses the corresponding genetic How does lactose induce the simpler to consider teria are first synthesis of the enzyme? It is perhaps bac- the inhibition of the synthesis. the synthesis of the enzyme can Why first then lactose necessary? Glucose has been used up in the phase of the bacterial development. but only after a negative phase that lasts around 30 minutes. The . metabolism continues terium has stocked. a repressor. [53] . Carbon and energy source. As long as this repressor is present. the result is the same. no ^-galactosidase first. Glucose is used up and the synthesis of /^-galactosidase starts. is is synthesized. and it starts synthesizing information.

no enzyme for its utilization is produced." This con- [54] .BIOLOGICAL ORDER this hypothesis implies that the repressor is is unstable and that its half-life short. the enzyme is useless. The growth rate of the constitutive bacteria is obviously lower than the growth rate of the adaptive ones. Anyhow. enzyme will not express itself if a repressor How does the repressor repress? Inductivity and con- stitutivity have to be considered anew. the bac- show that the constitutive bacteria contain about 30% more enzyme than the inductive ones. metabolite. In both cases. This the fact that part of the building blocks is is obviously due to utilized for the manulactose lactose facture of a useless enzyme. the When When enzyme is synthesized and the synthesis adjusted to the need. The structural gene carrying the information for the synthesis of a given present. insensitive to the whether or not lactose Estimates present. present. a preliminary attempt ization seems to be necessary. absent. When into a adaptive and constitutive bacteria are inoculated together of lactose. The effector of the regulatory mechanism is a repressor. is In the cases studied so far. is controlled by the "level" of this "Level" here means intracellular concentrawhether the essential metabolite is extrinsic or endogenous. the synthesis of the enzymes necessary for the synthesis of any given essential metabolites essential tion. Enzyme synthesis is of course limited by something. let us say a few minutes. in a medium devoid of lactose. Before any further discussion. Moreover. It at a general- might be stated that the synthesis of the enzymes required for the metabolism of the energy source is controlled by the over-all available energy sources. some others because is is the "operator" (explained in the next section) repressor. The sidase question will immediately be asked: How does /?-galacto- behave in a constitutive bacterium? Some bacteria are consti- tutive because they do not produce a repressor. is is Evolution has thus ended in a remarkable device. let us say by the availability of amino acids. inductivity "dominant. the constitutive bacteria are out- medium devoid grown by the adaptive ones and disappear. owing to natural selection. How does the repressor act? The is operator. terium contains yff-galactosidase.

desiraated by the letter The dominant the inductive + would have the o'' the constitution i+o+z + . and the constitutive gene i~. elaborate series of experiments has led to the conclusion that a when a bacterium mutates from recessive to dominant constitutivity. despite the presence of the inductive gene i + An hypothesis is that the gene produces repressor. The regulating gene produces — or does not produce —a specific repressor. no enzyme is formed in the absence of an inducer. the other i^z + In the absence . is dominant. Thanks to recombination experiments. specific gene is involved which has been called the operator and o. a The and the expression of the structural gene z+ of the chromosome i~z+ is blocked. 0+ gene being stitutive would be repressor. The func- tional unit formed by the operator gene and the structural gene has been called an "operon. . and next to it the regulating gene i. instead of being recessive.THE FUNCTIONAL ORDER elusion is two sets of based on the consideration of a heterozvgote containing chromosomes. The inductive gene i+ is said to be dominant. despite for constitutivity. The problem of the site of action of th. recessive (Figure 21). in position cis (Figure 22). The gene i+ imposes the inductive situation. It "sensitive" to the repressor. Thus. When constitutivity is dominant. that is. But affects only those genes located on the same chromosome. An extensive study has revealed that in some mutants constitutivity. the heterozygotes i + z + i~z+ manufacture the /3-galactosidase in the absence of inducer. of an inducer. The repressor produced thanks to the gene i+ acts on the "constitutive" chromosome carrying the constitutive gene i~ or on enzyme-forming system produced by the "constitutive" the chromosome.e repressor was solved by the discovery of a new category of mutants. despite the presence of the constitutive gene i~." The repressor formed by the combination of the aporepressor and [ 55 ] . no /?-galactosidase. The dominant cono"^ gene being "insensitive" to the was found it that the mutation o+ ^ affects not only (for the the gene z (for the /?-galactosidase) but also the gene v permease). i~o'^'z . no enzyme. The structural gene contains the informa- tion for the synthesis of the specific enzymes. the presence of the gene i~ responsible i+ is produced. one i + z+. Close to them is the operator gene o. it is known that the genes y and z are adjacent. .

dominant but only for the gene y+ located on the same The gene z+ is located on a chromosome carrying the wild-type operator o+ and is not active. is Inductivity dominant. The Operator Gene. No enzymes produced by the heterozygote in the absence of an inducer. located on the same both permease and iS-galactosidase are produced is in the absence of an inducer. 3. Constitutivity as o'' chromosome (position cis) . despite the presence of a repressor.BIOLOGICAL ORDER 1 - Inductivity i+ dominant operator + sensitive to repressor no/3 -golactosidase no permease 2ond3-Constitutivity operator 0^ insensitive I- dominant to repressor 2- 0'^/Z-*-y* in c/s fi -galactosidase present permease present 0VY+ no /3 inr/5 permease present 0<^/Z + in trans -galactosidase ^ •yj-galactosidase Opermease x repressor Figure 1. The operator are is of the wild type (o+) that is sensitive to the repressor. 21. i5e^ . are active: genes z+ and y+. The chromosome the constitutive operator o". Constitutivity as is dominant. 2.

for example.THE FUNCTIONAL ORDER of the corepressor would act on the operator gene. and enzyme synthesis is repressed. When the repressor is present. When no repressor is present. the operator gene is in position go. The structural genes z+ and v"^ "manufacture. the inducer. the operator gene o+ is in position go." respectively. Lactose. competes with the corepressor and prevents the formation of the repressor. When the inductive bacterium i+ is o + the operator is sensitive to the repressor. The Operon. When a repressor is present. and the question must be asked: How does the operator gene act? We have every reason to believe that the operator does not send messages to the cytoplasm. The repressor is produced from an aporepressor synthesized by the gene i+ and a corepressor that is a product of metabolism of the energy source — glucose. W' hen the repressor is absent. produce the enzyme-forming system. One hypothesis is that it acts by inducing a [57] . . the operator gene is in position stop. and the two structural genes z+ and v+ are "blocked" and do not OPERON regulating gene operator structural genes + Y+ ^____^ ~ ~ enzynrie forming systems /3-galactosidase permease co-repressor lactose metabolism Figure 22. /3-galactosidase and /3-galactoside permease only in the absence of the repressor. and the structural genes are active. There is a new problem now. the operator gene is in position stop.

however. All the members of a zymon work in harmony. is Co-ordinated control. if all the individual genes are linked and belong to the same operon. But the enzymes are there. and the activity of the already existing enzymes is depressed or stopped. the synthesis of the re- enzymes is repressed. but nothing more is known at the present time. A [58] . or first blocks. This corresponding to a right. for the genes of the is all tryptophan zymon. When the output ex- ceeds the consumption. And the activity of each enzyme depends on the activity of all the members of the zymon. This is zymon are some- the case. the genes zymon are not linked. It simple device. This would be the the genetic information for the zymon that is a unit. The study of bacteria has revealed the structural genes carrying the information corresponding to one times linked. Sometimes. acids. is When excess. the functioning of the enzyme of the biosynthetic path- way. Thus a and more sponsible mechanism controls the production of amino generally of essential metabolites. inhibition of enzymatic activity. The zymon is a functional whole. This conto exist in the arginine dition is known dual and the tryptophan zymons. One enzyme acts on the product of the activity of the preceding one. and if they continued to function. The most efficient and economical system of control would be a single system controlling the synthesis of case if all the members of the zymon. The other enzymes involved synthesis are activity of the in the particular chain of bio- consequently deprived of their substrate. Let us consider first the synthesis of an essential metabolite: tryptophan. happens that this accumulation is prevented by a very The end product of enzymatic activity inhibits. Preclusive and corrective feedback mechanisms. that the need. Refinement of regulation. is. the accumulation of the end product would continue. The group of enzymes that for the synthesis of a given essential metabolite or responsible more generally for a given sequence of reactions will be called a zymon.BIOLOGICAL ORDER functional change in the structural genes. the end product of a biosynthetic process a produced in repressor is formed which blocks the synthesis of the corresponding enzymes. But will be found. and the zymon is reduced or blocked (Figure 23). it And for this no explana- tion has yet been found. for example.

THE FUNCTIONAL ORDER OPERON structural genes reguloting gene operator L^^' enzyme forming systems Qd O/i Oy 6^ enzymes .

zymon. the repressor is one of enzymes different from the system submitted to the repression. The corrective feedback deals with the synthesis of an essential the end product of the metabolite. counteraction. 318-356. the bacterium. 26. 1961. no single molecule or group of molecules can be held responsible for *The reader interested in the control of protein synthesis and in the problem of regulation is referred to the excellent review of Jacob and Monod. or one group of persons. microorganism. one person. a repressor is the synthesis of ^-galactosidase. and to Jacob and Wollman. has been compared to a factory. [60] . The of all organism. In order to survive. it must adjust its enzymatic equipment according to the nature of its food and to the nature of its needs. And also. many more systems have to be analyzed before any attempt at a generalization can be made.* the tryptophan the end products of a system of Conclusion The study of enzyme synthesis has thus revealed that the cell is endowed with an elaborate dual mechanism which controls the activity and the synthesis of enzymes. The repressor prevents present. In the first case. Some in a readers may be tempted to ask the question: in a Who commands bacterium? Obviously. Things are different when the energy source glucose that is is is concerned. When produced. The organism has to cope with the variation of the environment. In a factory. Sexuality and the GeJietics of Bacteria. Academic Press. the corepressor is activity of the enzymatic system submitted to repression. In the other case. MoL BioL (1961). thanks to a corepressor a by-product of glucose metabolism.BIOLOGICAL ORDER negative feedback mechanism: in this case the control is corrective. the machine must be regulated in such a way that each one of the four nucleic bases and each one of the twenty amino acids is manufactured in just the right amount and proportion. The activity of the enzyme re- sponsible for the metabolism of one energy source prevents the synthesis of a system involved in the metabolism of another energy source. /. directs the activity the others. New York. the preclusive feedback with the metabolism of an energy source. Obviously. namely. or preclusive. but it is in fact a preventive. This mechanism could be considered as a feedback. to the last issue of the Cold Spring Harbor Syinposia on Quantitative Biology (1961). 3.

685-691. Angew.. eds. G.THE FUNCTIONAL ORDER the harmonious dynamic cellular balance. The enzymes and the comparator. A Syinposium on the Chertiical Basis of Develop7Hent. Repres(1958). Symposium on the Regidation of Cell Metabolism. Induktion. itself is metabolism the result of the interaction of and of the dual feedback mechanism. W. The Johns Hopkins Press. Wolstenholme. This formula should take on its full significance. it is simply suppressed. an essential part is function to compare the result of the \vork done by the machine with the information fed to the computer. ed. The functional order is the result of the interplay of the hereditary material. Structures and functions are the is complementary aspects of of order. sion.. New McElroy. There is no such thing as a comparator in a microorganism. E?izy??!es: U?iits of Biological Structure and Function (Henry Ford Hospital. Evolution has necessarily selected the fittest types. Biosynthese eines Enzyms. of the enzymes. Che?me. The living organism has been defined as an independent unit of integrated structures and functions. The organism a functional system REFERENCES Gaebler. When the work does not fit. of the metabolism. E. International Symposium). J. Boston. W. pany. Its food. Baltimore. and Glass. (1958). Instead. A now specific function can only be the expression of a specific structure. the struggle for life. Academic Press. H. Brown and Com- [61] . 11. eds. C. and O'Connor. Information. as proyided by the environment. O. D. (1959). M... B. each organism takes part in a compe- tition. life. The comparison a functional one. Ciba Foundation Little. Alonod. in which the functional values of the is different types are compared. York. (1956). is In a computer..

one discovers en- able to reproduce their kind is which are about 250 A in di- ameter. which to organize these enzymes a The ultimate unit of integrated structures and size. Two systems of order are then intertangled. Their volume therefore 8. functions must necessarily possess a minimal As matter of fact. They are strictly dies. the diameter of the smallest microorganism capable of independent reproduction When tities is about 5. The cell must at its disposal some 2. to grow. a living cell. as such. the is unable to metabolize. It must therefore have possess the genetic information for the synthesis of these in 2.000 enzyme species in order to secure energy and to synthesize all its building blocks. the cell/virus system survives and multiplies.VIRAL FUNCTIONS: ORDER AND DISORDER Introduction An kind. Viruses can develop only inside intracellular parasites. Viruses are able to reproduce their kind. These entities are viral particles. A cell infected by a virus very often Sometimes. The viral order is [62] .000 in the enzymes and some space proper organelles. tion organism is an orderly system of integrated its structures and functions able to metabolize and to reproduce The microorganism or the cell is the ultimate unit of integra- and reproduction. however.000 times smaller than that of the smallest microorganism. viral particle.000 A. Yet. Ulthuate here means smallest. one studies the living \vorld. Viral multiplication thus poses a certain number of problems. however. They contain proteins and nucleic acid and exhibit a constant and welldefined structure. and to undergo binary fusion.

viral functions. which means a unit of virus. the poliovirus. Various aspects of viral order. will be taken as a proa virion. The virion is a polyhedron. modern. Protein coat Subunit Nucleic acid Figure 24. and to be terminology. It has been crystallized and its crystals studied by X-ray diffraction. The most general idea of a viral particle can be derived from the study of one of the simplest and smallest viruses. It is composed of a protein shell en- closing the nucleic acid (Figure 24). Schematic Representation of a Viral Infectious Particle (Virion). The virus of poliomyelitis. and cell/virus interactions will be discussed in this chapter. In order to simplify things. or cancer when the mammalian cell is involved. As a consequence. the infectious viral particle will be called virion. made of subunits. the cellular order is perturbed. \^iRAL order: the infectious viral particle. X-ray diffraction shows that [63] . according to recent to spare words. or virion. arranged in an orderly fashion. totype of This virus has been obtained in a high degree of homogeneity and its chemical constitution established. the capsomeres. The The genetic material (nucleic acid) capsid is is folded and enclosed in a coat or capsid.VIRAL FUNCTIONS: ORDER AND DISORDER superimposed on the cellular order. and the results may be as different as "lysogeny" when bacteria are concerned.

the virion is metabolically inert. acids. its The is. viral genetic material. These conditions exist only inside a cell.600 amino acids. Even if some of the proteins are endowed with a potential enzymatic activity. the maximum number of amino acids that can be organized in the viral nucleic acid is 1. is number of protein It species.700. which is the critical living mass. No is other "biological" particle or a large organelle yet known which composed of number of asymmetrical subunits exhibiting relations of symmetry. This type of structure seems to be able to form only a viral coat and nothing else. built of subunits. that itself. to synthesize the specific viral proteins and must necessarily be in an environment where building blocks and energy are provided. The in structure of the viral it infectious particle is extraordinary the sense that is is unique.600 amino of smaller molecules. The molecular weight of of the virion is approximately 6. It is built up out of 60 structurally equivalent asymmetric units of approximately 60 A diameter. the conclusion was reached that a sequence of three nucleotides was probably necessary to take care of one amino acid. in order to express potentialities. The capsid can only be and is in fact built of subunits. This type of structure is clearly seen in electromicroscopic photographs of many viral species. The genetic material of the virion therefore cannot directly build a molecule composed of 49. The nucleic acid is composed of about 5. which the protein represents about 62%. is but it can build a number Each capsomere in fact made up of approxi- mately 620 amino acids. Therefore. The protein coat of viruses or capsid close contact in a form a is actually made up of subunits or capsomeres. to duplicate The essential function of the protein viral coat is to provide a cell to cell means for the transmission of the genetic material from [64] . they roughly spherical shell of about 300 A diameter.200 nucleotides. An entity containing a small five. cannot secure energy and cannot synthesize its building blocks. If 60 protein subunits of that size are packed together in way consistent with icosahedral symmetry.BIOLOGICAL ORDER each virion possesses an icosahedral symmetry. In the discussion of the code. the nucleic acid 28%.7 X lO^*. let us say one to unable to do much work. The protein of the capsid contains about 49.

let us say poliovirus. it is The proof of the effectiveness of the infection the production virus is by the infected cell of a new generation of virions. The genetic material is RNA in some viruses.000 particles. and which is an essential and clear also that the genetic material of information (a) for the synthesis of the pro- teins of the viral coat. This is the so-called vegetative phase of the viral are not life cycle. A virion never contains both nucleic acids. the synthesis of cellular structures a well-ordered process. properly handled and not altered. viral proteins are synthesized. subject to a regulating system? If a viral regulating system does exist. Moreover. reproduction from the genetic material only specific feature of viruses. They -appear around the third hour. a An elaborate system of repression and induction based on feedback mechanism controls the balanced as synthesis of enzymes. and (b) for autonomous reproduction. pathological particles of Are viruses. and DNA. in others. Every every micro- RNA nor a organism without exception. Remarks on the control of viral functions. Approximately 6 to 8 hours after the infection each cell contains about viral 100. how do viral and cellular functions interact? The study of bacteriophage should allow us to answer these questions. It a virus has to possess the is is something which neither a cell microorganism can accomplish. The nucleic acid of a If number of viruses has been if extracted and purified. infective particles. [65] . the cell damaged and dis- integrates. In a normal. DNA cell. and the genetic material is multiplied. infect cells. among which only about 500 is are infectious. reproduced from its genetic material only. finally As a result of viral multiplication.VIRAL FUNCTIONS: ORDER and a protection against the adverse conditions AND DISORDER in the encountered outer world. An organized infectious entity possessing only one type of nucleic acid can only be a virus. The may The virus. like normal cellular constituents. and thereafter will increase in number. produced during the first phase. When a cell is infected by a virus. At the is cellular level. noninfected is cell. a viral infection the penetration into a cell of the genetic material of a virus. Mrions. possesses both types of nucleic acid. Viruses are sometimes considered cellular origin.

the bacteriophage becomes external attached to the bacterium (Figure 26). But when the tip of its tail into contact with the bacterial wall. (a) a contractile outer sheath. composed of is a ball of DNA surrounded by is a protein coat. Figure 25. tail The is contracts. is The synthesis of phage protein initiated. (b) an internal tube. The bacteriophage is more complicated than the poliovirus. and (c) tip. The genetic material through the tube into the bacterial cytoplasm. It has a so-called "head" and a so-called "tail" (Figure 25). The head is a hexagonal particle prism. The internal tube penetrates through injected membrane a inside the bacterium. and 5 minutes after the [66] . The tail a long cylindrical structure. Schematic Representation of a Bacteriophage Particle. sheath of the the An enzyme present at the extremity of the phage tip attacks the bacterial wall. The comes bacteriophage is not motile.BIOLOGICAL ORDER Bacteriophage Life-cycle of a virulent bacteriophage. which composed of a. All within minute. this happens Then the so-called vegetative phase takes place.

After 12 to 20 minutes. The bacterium is lysed and liberates some hundred [67] . The chromosome c is disintegrating. the is tail appears \yhere it should appear. bacterial chromosome has disappeared. 26. This endolysine depolymerizes the mucopolysaccharides of the bacterial wall.VIRAL FUNCTIONS: ORDER infection the genetic material AND DISORDER of the bacteriophage starts multi- plying. and the autonomous multiplication of the phage genetic material is initiated. has injected genetic material into the bacterium. The bacterial 3. a peculiar phage protein has been produced. (b) 2. condensed into and the subunits of the head are assembled around the ball is DNA <v/w» — genetic material of the bacteriophage endolysine • phage proteins Figure 1. The \vill infectious particle formed. Finally. Its building blocks (nucleic bases) will be incorporated in the phage genetic material. and phage particles are formed. A lytic enzyme (endolysine) bacterial wall will be will be vegetative bacteriophage. thus liberating the bacteriophage particles. The it Life Cycle of a Bacteriophage. by a process that is not understood. 4. of the In the meantime. The produced by the hydrolyzed and the bac- terium will l)-se. The proteins are organized around the folded genetic material. Beginning of the vegetative phase: Phage proteins are produced. the be localized at the tip endolysine. the bacteriophage particle has attached onto the receptive its Infection: (a) bacterium. of nucleic acid. More phage material has been synthesized. the phage a ball. \yhich in the particle tail.

BIOLOGICAL ORDER bacteriophage particles. called viriile?it phages. called temperate phages. Others. Some bacteriophages. are multiplied only by the lytic cycle. The so-called "lytic cycle" of the bacterio- phage has been completed. may evolve in two directions. / c .

The lysogenic bacterium multiplies a specific structure: the prophage perpetuate to (etymologically. immunological specificity. shape. 1953). etc. the bacterium grows and divides. it dies. CDnon-lysogenic b ( [ >*/ Productive infectio i :'c^ phase ^g^ Vegetative [^~:cc:<-J CHD >{Em^ Figure 28. which might differ by a number size. A number of experimental data have led to three essential con- clusions (Figure 28). When this potentiality is not expressed. it will perpetuate the potentiality to produce B. [69] . of properties such as host range. Diagrammatic Representation of Lysogeny (after Lwoff. When infected with B. us say A or B. "before the phage"). A given bacterial species can be infected by let a of different strains of temperate bacteriophages. When it is expressed. The number prophage.VIRAL FUNCTIONS: ORDER possesses AND DISORDER and perpetuates the potentiality of producing bacteriophage particles in the absence of infection. A given bacterium infected and lysogenized by the potentiality a temperate bacteriophage A will produce bacteriophage A. The bacteria that perpetuate the possibility of producing bacteriophage in the absence of infection are called lysoge7iic.

itself after an infection. How is it possible that one is dual behavior? This But there prophage. and the vegetative phase culminates with the organization of infectious particles. It carries the information for the production of bacteriophage particles. according to the bacterial species. is always attached to the same site. is homologous chromosomes takes place. [70] . the genetic material of the bacteriophage bacterial rial attached to the chromosome. 2. 3. originated from the Prophage and vegetative phage. the its chromosome 2 multiplies at own pace. of the bacterial chromosome. has chromosome. bacterial in favor of the hypothesis according to which the genetic material of the phage has. Cytologists know that during a certain phase of the maturation of the sexual cells. behaves differently from the vegetative genetic material. the bacterium is lysogenic. The prophage that is. The nature of unknown. every 20 to 50 minutes. viral During the vegetative phase. admitted that the attachment of the of the bacterial viral genetic material on a specific site the consequence of a correspondence of structure of the chromosome two is interacting nucleic acids: the bacterial one and the viral one. or locus. duplicates itself alongside the bacterial chromosome. When. In a lysogenic bacterium. The phage does not express its potentialities: no viral proteins are to be found in a lysogenic bacterium and of course no infectious particles.BIOLOGICAL ORDER 1. etc. the prophage. During chromosome is autonomous. That a given prophage site is is able to reach and recog- nize a specific chromosomal something strange. The prophage There is is the genetic material of the bacteriophage. it is. The genetic mate- of the bacteriophage. long ago. a pairing of stick The two chromosomes regions. This correspondence. the phage medium. as prophage. The receptor. temperature. another difference between vegetative phage and vegetative viral The chromosome expresses its potenpro- viral proteins are synthesized. It duplicates itself every or 3 minutes. nature of the the vegetative phase. A given prophage only one prophage per bacterial chromosome. tialities: is and the same structure exhibits such a one of our problems. submitted to bacterial control. whether it is homologous or complementary. to each other at their homologous is the forces that intervene in the process it is In any event.

however. the lysogenic bacterium does produce phage. the answer. But thanks to the presence of the prophage the lysogenic bacterium exhibits another property. the genetic material of phage A does not multiply and of bacteriophage. and each one has liberated some [71] . These strains are called inducible. In other can be induced at will in practically all of the population. Other types us say B.to 10~" Induction. is diluted out in the course of bacterial is multiplication (Figure 29).VIRAL FUNCTIONS: ORDER AND DISORDER defined as a bacterium that perpetuates produce bacteriophage in the absence of infection. spontaneous because \e do not know cause. Then within 10 minutes. not genetically related to bacterioif prophage A were not cytoplasm of the lysogenic bacterium carr\ing prophage A which prevents specifically the vegetative multiplication of the genetic material of phage A. the production of bacteriophage cannot be modified. The The simplest hypothesis study of lysogeny should provide is that viral functions are not expressed in a lysogenic bacterium because they are blocked by a repressor. ImiMumty. Let us assume that our bacterium has been lysogenized by a temperate bacteriophage A and therefore perpetuates a prophage A. The per bacterium per generation. Viral functions are not expressed. Phage proteins are not produced. the so-called im- munity. The superinfection by the homologous bacteriophage A does not result in a vegetative phase. Moreover. From time to time. frequency of spontaneous production In some lysogenic strains. the is the so-called spontaneous its phage production. phage A present. This strains. something can multiply vegetatively. All the bacteria have been lysed. Thus. but behaves as an inert particle. The genetic material of the infecting it bacteriophage A is injected into the lysogenic bacterium. Inducible lysogenic bacteria are irradiated with a suitable dose of ultraviolet light. the bacterial population disappears. a lysogenic bacterium perpetuates the potentiality to produce phage. The bacteria continue to grow for about 45 minutes. This let immunity specific. probability that a lysogenic population will produce infectious particles varies with the strains from 10 ~. and we have to know why viral functions are sometimes the A lysogenic bacterium was power to expressed and sometimes not. just as must be present in the Here again a specific repressor of viral functions can be visualized.

and One of the daughter bacteria (3) contains only the prophage. by an irradiation with ultraviolet The vegetative phase of both prophages and the super- infecting phage are initiated. This lives in is harmony with its prophage. most extraordinary. The problem of the control of viral development is thus posed in the most dramatic way. of i V y- (p) or corresponding autonomous D. A bacterium The prophage behaves as if it suddenly. the lysogenic bacterium has dies. gous phage. 4-5. for example. the vegetative phase of the bacteriophage initiated.N. is induced.BIOLOGICAL ORDER 100 infectious particles. A. as a result of an irradiis were a normal bacterial gene. The genetic material of the superinfecting phage does not initiate is the vegetative phase. 29. lysogenic bacterium produces bacteriophage and to the irradiation. does not divide. previous been infected with a mutant of the homologous phage toward which the bacterium is immune. The superinfected bacterium light. and the i^r prophage D. the mutant will develop together with the prophage. is The lysogenic bacterium superinfected with a mutant of the homolodiluted out at each division. homologous superinfecting phage Figure 1-3. [72] . And ation. A. Immunity.N. If. and infectious particles of both types will be produced.

This These data are consistent with the hypothesis that in the lysogenic male bacterium the expression of the viral functions is blocked by a cytoplasmic repressor. an inducible prophage penetrates zygotic indnctiov. All these agents are known to be mutagenic and oncogenic. the vegetative phase is initiated. the vegetative phase initiated. epoxides. Ultraviolet not the only inducer. and viral functions are How is could the repressor act? The antibiotic chlor- amphenicol known to block the synthesis of proteins. that is. The prophage A of Escherichia coli K12 is located on the bacterial chromosome close to one of the genes controlling the utilization of galactose. induction of a lysogenic bacterium is not homologous to a mutation. also inducers light has AND DISORDER is an inducing effect but A number of physical or chemical agents are — among them. cancer-inducing. If only the female if a lysogenic. a fact that is both male and female are lysogenic. female is lysed. Anyhow. phage development is induced when. If bacterial [73] . and ethylenimines.VIRAL FUNCTIONS: ORDER Inducing agents. nitrogen mustard. the level of the repressor decreases is below a given threshold. nothing happens either. As a consequence. Thus. the theory that the inducing agents depress or block the synthesis of the repressor. and the is genic male has penetrated into the nonlysogenic female. as a result of conjugation or zygosis. repression ceases because the cyto- plasm of the female expressed. is devoid of repressor. If conjugation. things are quite different (Figure 30). or ^

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hich has a greater affinity for the repressor. The fact that inducing agents are mutagenic is not a great help: organic peroxides. But the fact that inducing agents are able to initiate malignant growth Perhaps induction will help us to understand is rather exciting. cancer. X rays. which enters into the female at about the 25th minute after conjugation. gamma rays. or a more stable repressor. a A tures noninducible prophage could be prophage which manufaclater. but the reverse is is certainlv not true. This problem will be discussed Zygotic induction. But a lysogenic male mates with nonlysogenic female. Since the prophage is no longer repressed. as the As soon prophage attached to the chromosome of the lysobacteriophage particles are produced. more repressor. nothing happens after is worth while noticing. When the prophage of the lysogenic male enters the nonlysogenic female. into a nonlysogenic cytoplasm.

Prophage during Bacterial Recombination. If this interpretation correct. zygotic induction. The vegetative phase is not initiated. — phage proteins X repressor Figure 30. Upper row: The lysogenic male (1) (repressor present) injects its chromosome and prophage into the lysogenic female (2) (repressor present). It could be that the viral function necessary for the onset of the is vegetative phase is the synthesis of a protein. it should be applied to infection and lysogenization. zygotic induction does not take place in a large fraction of the zygotes.BIOLOGICAL ORDER phage D.N. [74] . The vegetative initiated: this conjugation takes place in the presence of chloramphenicol.A. Lower row: The some and prophage phase is lysogenic male (4) in a is (repressor present) injects its chromo- nonlysogenic female (no repressor).

and the bacterium viral now lysogenic. It continues to we shall try to understand why produce the repressor. Something irreversible has happened.) acts in favor of lysogenization (Figure 31). If a protein is is produced is the vegetative phase initiated. the viral DNA duplicated. Thus. But.VIRAL FUNCTIONS: ORDER AND DISORDER fected Infection and lysogenization. does. namely. like any other genetic material. It is now a prophage. the viral site. The outcome of the infecon the genetic constitution of the temperate bacteriophage. material of a bacteriophage. DNA The it is to this receptor later on. If a repressor is produced first. is a repressor. A nonlvsogenic bacterium inby a temperate phage may either produce infectious particles and be Ivsed.) a low temperature (20°C. the expression of the viral potentialities is blocked. A high whereas temperature (40°C. A mutant ind^ was discovered which had lost its original inducibility [75] . and finally infectious particles are produced. and the process irreversible from More proteins are synthesized. and the fate of the bacterium. the behavior of tion depends the bacterium/phage system. favors the establishment of the lytic cycle. or become Ivsogenic. thanks to the correspond- DNA ence of structure with attaches itself a given chromosomal locus. After the seventh minute the decision made and cannot be modified. But with a given temperate bacteriophage. The genetic mutants. and it will be reinforced by the study of Mutations affecting inducibility. and the autonomous cytoplasmic viral is unable to duplicate itself. the experimental data point toward the conclusion that the response of the infected bacterium depends on a primary determining event. What It is this event? that antibiotics was found first which block protein synthesis. The low or high temperature acts only during the first 7 is minutes after infection. are controlled by extrinsic factors. may undergo muta- The wild-type A ind+ is inducible with ultraviolet light. The fraction of the infected bacteria that will be lysogenized varies effectively from less than \^c to more than 99%. such as is chloramphenicol. something then on. else. favor the lysogenic response. that the The a hypothesis event can be the synthesis either of protein or of first. tions. The repressor hypothesis needs of course to be reinforced. Proteins cannot be synthesized. prophage multiplies.

and infectious particles are produced. ^ phage proteins X repressor Figure 31. is.) in the presence of chloramphenicol.V. or in a bacterium irradiated with ultraviolet rays. The vegetative phase is initiated. The Fate of the Infected Bacterium.). ble with ultraviolet light Double lysogenic strains The mutant not inducihowever. and the genetic material of the phage reaches its chromosomal locus and becomes a prophage. a As with ultraviolet light. can be obtained that carry two prophages A: [76] . sensitive to zygotic indu<iition. although all other characters of the mutant were the same as those of the original strain. is The nonlysogenic bacterium 1-3: at infected: low temperature (20°C. X X •\AAAphQge A.BIOLOGICAL ORDER nfection 20°. 4-5: at high temperature (40°C. The first activity of the genetic material of the phage is the synthesis of a protein. or in first bacteria deprived of food. U. N. the vegetative phase is not initiated. starvation 40°. Chloromycetin. the repressor. X ^. activity of the genetic material of the consequence the synthesis of phage protein is repressed. lysogenizotion ' vegetative phase X X X ^ ^ X X X X V X ^ X X X X X X X y X X X D.

When a bacterium carrying prophage A ind+ is irradiated with ultraviolet light. Table V. Inducible .VIRAL FUNCTIONS: ORDER the inducible one. The noninductive mutation A ind" is dominant. the phage A ind+ cannot be induced. AND DISORDER these strains are When none of the bacteriophages develop. Variation of Inducibilitv. irradiated with ultraviolet light. it is induced and will produce phage (Table V). In the presence of A ind". and the noninducible one.

Ability to lysogenize . Mutations Affecting the Life Cycle.BIOLOGICAL ORDER Table VI.

and the is genetic material of the phage. Defective Prophages. . more of these proteins is impossible. And apparently.VIRAL FUNCTIONS: ORDER AND DISORDER Some are able to the synthesis of one or manufacture the proteins of the phage. is not multiplied. It seems as though the vegetative phase could be characterized by a sequence of events. In others. so far as known. the chromosome of the defective bacteriophage Table VII. in the absence of protein synthesis. Each event has to be completed so that the following one can be initiated.

Moreover. This happens also if the repressor balance is upset by inducing agents such as ultraviolet light. 1. The most important aspect of the reproduction of the viral genetic material has not yet been discussed. and they produce the proteins necessary for the unable to produce onset of the vegetative phase of their It own life cycle. has been admitted that the prophage does not enter the vegeta- tive phase If this because it is unable to produce some specific protein. Their properties seem to be controlled by their position in the bacterium. 2. the homologous prophage enters the vegetative phase. Let me repeat that the virulent mutants of a temperate phage are a repressor and are insensitive to the repressor produced by the original prophage. The synthesis of this protein may be blocked by a specific repressor. If a lysogenic bacterium is infected with a virulent phage genetically related to the prophage.BIOLOGICAL ORDER the autonomous if replication of the viral genetic material can take is place only the genetic material of the phage no longer exposed to the repressor. the behavior of two structurally identical viral chromosomes might be different. the virulent mutant. whereas the pro- phage multiplies. the chromosome of the superinfecting bacteriophage does not multiply. All the experi- mental data are thus in agreement with the hypothesis that autono- mous multiplication of a phage requires the manufacture of a protein. This happens when the prophage of the lysogenic male enters a nonlysogenic female where no repressor is present. which does multiply and which does produce these proteins. should induce the vegetative development of the prophage. In a lysogenic bacterium. the prophage is duplicated in harmony with the [80] . They develop despite the presence of a repressor. whether it is attached to the chromosome or to an episome such as the sexual F factor (Figure 32). Thus. hypothesis corresponds to the facts. Position is the fourth dimension of the prophage. Interactions of Cellular and Viral Functions Cellular control of viral reproduction.

The prophage is obviously subject to the system that controls the duplication of the bacterial chromoquestion some. is: What is this system? As a matter of exist? fact. that is. occurs every 20 to 50 minutes. the first Does such a system .VIRAL FUNCTIONS: ORDER cellular organelle to AND DISORDER one duplication which it is attached.

In one case. The enzymes for tryptophan synthesis may be produced while those for methionine synthesis are repressed. It happens that this gene galactoA. Thus the genetic material of a virus. A part of the receptor of prophage A the gene that carries the information for galactokinase (the kinase can be exchanged with a part of phage kinase is enzyme that phosphorylates galactose). the prophage behaves as In the other case. The prophage behaves is were a bacterial element. all The existence of such functions chromosome can only be postu- strong argument in favor of this hypothesis is that in a cell chromosomes divide not only at the right time but also at the same time. submitted to the cellular system of control. Obviously. The study of the behavior of the defective prophage after induc- tion has revealed the existence of a set of functions controlling the duplication of the viral nucleic acid. The reverse is also true: is cellular gene may be under viral command. of the induction and repression of enzyme synthesis. In a bacterium. some unknown factor commands the the duplication of the chromosomes. has revealed a different situation: the various individual systems controlling enzyme synthesis are inde- pendent in the sense that one may be blocked while the others are working. when a attached to a cellular organelle.BIOLOGICAL ORDER and persist in a prophage which is reproduced as nucleic acid and which can be perpetuated without expressing its potentialities. the A chromosome and as if it the prophage act as a whole. were a viral gene. nonlysogenic If Sahnojiella produces given somatic antigen lysogenized [82] . the gene galactokinase multiplies too. Are viruses able to take command of cellular functions? The reader realizes of course that the question would not be posed of a if a positive answer could not be provided. Viral control cellular functions. given chromosome. a bacterial gene behaves as This probably means that viral. A 10. and when this The gene galactophage A multiplies vegetatively. whether cellular or if it if it and the "foreign" nucleic acid it may carry constitute a unit and behave as a whole. The prophage the bacterial attached to a specific receptor site of is chromosome. a were a bacterial gene. so far as replication is concerned. now part of phage A. for the duplication of the cellular lated. The study of the functional order.

Thus a virus may be subject to the control system of a bacterium. and not in position trans (when they are in different structures). both for replication and for expression. turn produces another antigen The mechanism by which is proa phage 15 controls the synthesis of a bacterial antigen still mvsterv. Thus may be multiplied as genetic [83] . A much more fascinating situation has been disclosed A short digression necessary here. Things happen here as if the bacterial gene were obeying some viral mechanism of control. The gene is with galactogalactokinase and the prophage A are closely linked. dies. either as a result of an infection or as a consequence of an induction. then the enzyme galactokinase is synthesized in large amounts in the absence of an inducer. Strictly speaking.VIRAL FUNCTIONS: ORDER with given phage AND DISORDER a e 15. If the partners are properly selected. the defective prophage is derepressed. a large variety of heterozygotes may be obtained. This synthesis takes place only when the gene and the phage are in position cis (when they are part of the same structure). When a defective lysogenic bacterium is induced. the disease is not viral. but the bacterium nevertheless result of an infection. and the hypothesis that the genetic material of viruses has is evolved from cellular genes currently admitted. bacteria contain only a small synthesis of the amount of A is marked present. Both may be attached to the sexual factor F. A few hereditary. The defective prophage possesses some of the normal viral genes but behaves as a potential lethal gene (Table VIII). enzyme takes place only when an inducer But when phage A develops vegetatively. nonviral diseases of bacteria are known which are controlled not by a defective prophage but by genetic structures that have to be visualized potentially lethal genes. and a bacterial gene to the control system of a virus. Viral and noxviral diseases. as bacterial genes. and in 15. it stops producing the antigen 10. and thus transferred from the male to the female. The disease is not the and infectious particles have not been formed. It happens that in the absence of an inducer the galactokinase. perhaps a viral operator. An abnormal vegetative phase is initiated. Bacteriophage particles are not produced. diseases. kinase. a virus Viruses and cancer. Between these hereditary series of diseases and viral an almost complete intermediary steps has been disclosed.

BIOLOGICAL ORDER Table VIII. Inductive gene . Genes and Viruses.

But the question possible It is is asked: is What can originate from order? the only answer disorder. viral functions are repressed. the nature of the cellular modidue to the virus or provirus and responsible for malignancy is still unknown. must not cell the cell it infects. When may endowed with its the potentiality for producing virus infection. Yet it is interesting to consider the animal cell/provirus system. The cell original cell was normal. according to the nature of the one way could be cell and of the virus. in a order to be oncogenic.VIRAL FUNCTIONS: ORDER for AND DISORDER it cancer. cell it is is sometimes killed. It or the other. The cancer problem fication not solved so far. the cell modified by the virus. the bacterium is an is which is organism. clear that the superposition of two types of structural order may be responsible for functional disorder. something more: the animal cell/virus system is it multiplies and kills its now host. as a result of an alteration of the cell's environment. animal cell/oncogenic virus systems. these The transformed yet-unknown factors. The it infected survives. Biological order origin in primitive disorder. Whether cell has lost its sensitivity to or not the genetic material of the virus multiplies autonomously as provirus is not yet known. Darkness comes from had its light. not the oncogenic virus fact that specific repressors are a that kills the cancerous animal but the cell/virus system. Injected into an animal. As already stated. At least one thing is certain. and kill killed. cold from heat. When. But there is malignant. it is immune also be modified in morphology and physiology. low virulence for the It is. According to the Platonic concept. to control The known some viral functions opens the problem of oncogenic viruses. subject to the factors that control reproduction. It in the absence of infection. Structures and functions [85] . new way of approach to all things originate in their opposites. pleasure from if pain. The animal as if it were independent. may mean a high degree of pathoitself genicity for the animal. however. sometimes to super- survives. A And virus. In some of the Viral antigens and infectious particles are not produced. viral antigens and viral particles are produced. viral functions are derepressed. the normal cell a malignant the host is cell behaves in the dependent part of an organism.

Genetic control of viral functions. Burnet. VVollman. 1709-1714. Lysogeny. S. — Biochemical. Interscience Publishers. New York. The Viruses J. 92-111. eds. 1-39. E. E.. transduction and cancer genesis. viral diseases Lwoff. F. M. and Stanley. Control and interrelations of metabolic and of bacteria. (1958-1959). La sexualite des bacteries. F. New York. Masson. series 54. and Jacob.BIOLOGICAL ORDER are complementary in processes. New York. L. The Harvey Lectures. T. Monographies de rinstitut Pasteur.. F. University of Texas Press. Nature. (1954-1955). John Wiley & Sons. [86] . New York. Volume I. Press. Luria. Bacteriophages. pp. Paris. A. Biological disorder logical order. series 50. (1959). H. (1959). F. Biological. New York. and Jacob. Austin. WoUman. M. Academic and Klug. Academic Press. Academic Press.. pp. and Biophysical Properties. normal processes as well as in pathological is the complementary aspect of bio- REFERENCES Adams. AL. Jacob. Structure of poHomyehtis virus. Finch. 43-59. A. The Harvey Lectures. 183. General Virology.. W. (1959). L. (1959). (1959). 43-59. In Genetics and Cancer. (1953). E.

BIOLOGICAL ORDER AND ENTROPY Sources of Energy a An organism is a maciiine. When is the organism possesses chlorophyll. The organism performs oxidoreductions. or mineral sulfide. Max Delbriick has described an organism as a system of flux equilibrium which takes matter and energy from its environment. It is not coal. the reaction AH2 This is + /z Oo —> A + HoO respiration. The organism. may be written: AHo hydrogen donor (reduced If. The general equation as sugars or amino acids. the source of energy energy in this is light. case. for example coal. When the organism is devoid of chlorophyll. but the combustion The living machine. however. in order to operate. that causes the steam engine to operate. of coal. Aietabolism and syntheses consist essentially in the mobilization of the energy of the chemical bonds of the food. burns more-or-less complex substances: organic com- pounds such as hydrogen compounds such carbon monoxide. machine has to be fed something. It needs energy. the provided by the chemical bonds of the food. the hydrogen acceptor oxygen. -f B hydrogen acceptor (oxidized state) — is A oxidized + BHo reduced acceptor is state) donor for example. or even hydrogen. In order to operate. must also be fed. [87] .

and the free energy of reactions ( measure of the maximum amount of work — obtainable. In the reaction: .BIOLOGICAL ORDER Equations In any chemical reaction. one has to distinguish between the total heat of reactions ( — aH) a The free energy F gives aF).

" that is. the probability increases. namely. Boltzmann constant and D is the measure of the The state of rest which an isolated system tends to reach corresponds to a maximum entropy. and an increase of order to a decrease of entropy.38 X 10" ^'^ and P the number corre- "elementary complexions. Entrop

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is a measure of atomic disorder. to the jumps of the atomic system from one metastable structure to another. An increase of disorder corresponds to an increase of entropy. that 1. from a thermodynamical viewpoint." "Elementary complexions" spond to the "discrete configurations. Entropy = k log D where k is the atomic disorder. Entropy Physicists know that every isolated system changes of rest. being a measure of disorder. is The between entropy and probability given by the Boltzmann- Planck formula: Swhere S of is k In P the entropy. The entropic a measure of a difference is The value of entropy between an initial and a given by the formula: final state. But during reproduction two organisms have been pro- duced out of one. to a maximum Now probable relation entropy. The measure of entropy is radation energy. In a from a less probable to more state. in such is a a way state as to approach of a definite state It This a state of rest of equilibrium. and so does entropy. [89] . k the Boltzmann constant expressed in is. we have to consider the production or reproduction of order in its most general aspect. Order has increased.BIOLOGICAL ORDER AND ENTROPY Reproduction as well as maintenance involves a degradation of energy. that is. disorder. closed system evolving is related to probaa bility. corresponds not to loss but to a deg- change during such an irreversible process is the difference between the entropy at the end and the entropy at the beginning. As our subject is biological order. ergs per degree centigrade.

A living system cannot be closed. as a result of reproducentropy has decreased. How can we measure the atomic order. the negative entropy in a living organism? [90] . This negative entropy. And we may D is a a measure = k log D. If entropy S /flogl/D." represents the quality or grade if admit with Schrodinger that measure of atomic disorder. its reciprocal \/D will be of atomic order. the decrease of entropy corresponding to the increase of order.BIOLOGICAL ORDER Thus. the degree of atomic order or the availability for work. As already stated. should be clear that the live the organism plus a reasonable — system under consideration is that is. we have to measure corresponds to tion. nucleic bases. Has the over-all entropy of the system represented in the equation for metabolism and reproduction increased or decreased? To answer the question. Let us now consider the equation of reproduction: (Organism) 1 + food —> As (organism) 2 its + waste + heat The organism has synthesized building blocks — its essential metabolites. nor a fortiori isolated. negative entropy or negentropy N= Life and Entropy A An is system is said to be closed its when it does not carry on exchanges closed system is of matter with isolated surroundings. "negentropy. unlimited — supply of food. like the systems studied It by physicists. Thus. that is. A said to be when it does not interact in any its way with its surroundings. die. and amino acids — and has organized them into specific macromolecules. organism separated from surroundings will soon and therefore an abstraction. Physicists have found it convenient to have an expression for its reverse. living a result of the functioning of the machine. that is. Increase of order a decrease of entropy. of energy. entropy is a measure of atomic disorder. work has been produced and energy has been degraded. order has increased.

BIOLOGICAL ORDER AND ENTROPY Information and Negentropy This brings us to the negentropy principle of information. information is measured in entropy units: /i = 1. of any specific organization can thus be expressed For example. The information / is /i = K In Po based on scarcity. The /o = with Pq probable or possible outcomes. In 1929. This definition of information is The lower the probability. the higher the scarcity and the higher the information. And the demonstration has been provided that information corresponds to negative entropy. When number of possible answers when no some information is gained. the negentropy of a telephone net- work with 108 subscribers can be calculated to be 4 X 10""^. The similarity between the two formulas of entropy S = k\nP and information /i is = K In Po is obvious. that is to say. If the Boltzmann constant used. then is number of possible answers reduced.38 X 10-i«lnPo The information in entropy units. Leo Szilard discovered the existence of a connection between information and entropy. Information initial situation is a function of the ratio of possible answers before and after. One considers a problem involving information the tion" is a certain available. is What information? We shall follow Brillouin's analysis. [91] . one single outcome selected. The final situation is with Pi = 1. and "complete informais means only one possible answer.

organized in number a given sequence. we can calculate the of possible arrangements of the 10" units. This organization should perhaps be considered also in the calculation of the amount of information. or a nucleic acid. If we apply the formula /= it 1. But the problem of [92] . It contains a few thousand enzymes and a large number of other specific macromolecules. visualize the structural is a protein. Since this is so.BIOLOGICAL ORDER What This is. is the One could consider that the negentropy of an organism sum of the negentropy of its specific macromolecules. the negentropy. and that today there is only one. In the bacterium Escherichia coli. we have assumed that at the origin there were 4^0" possible outcomes. But an organism is much more than its genetic material. But organization of in all the problem can be simplified. terial is composed of 10" nucleic bases belonging to 4 categories which is the hereditary mamaterial. is the information. that is.38 X X 10-16 InPo follows that /= 1. The origin the macromolecules of an organism has its the organization of the hereditary nucleic acid. selected from among a large number of probabilities. of an organism? as pointed out by Brillouin in his classical book. explored field The Negentropy Physicists of Biological Organization know that in a machine the structural negentropy represents the information or organization of the machine. that is. ergs/°C. an unof research. This figure was obtained by considering only the genetic material. It is 4^^'. How can cell? we negentropy of an organism or of a Each species of macromolecule.38 10-16 In 410' The structural negentropy of the bacterium Escherichia coli is around 2 X 10 -^ entropy units. For the calculation of the information of our bacterium. part of the organization of the living machine. whether it is a polysaccharide.

radical development and a broad extension of the notion of entropy and to something quite different from the classical physical concept. its information will be destroyed. that this cannot be measured in terms of entropy units. functional coupling between the active units of a into account in the calculation of And Linschitz has proposed that this type of organization should also be taken what he calls a "biological I negenlike to tropy. one given molecule of guanine in a given gene is replaced by a molecule of adenine. This corresponds to a improbable system but a fitted for certain functions. one nucleic base of the genetic material sometimes replaced by another. The study cell. of the functional order has revealed a very high degree of nonstructural. original. for example. This alteration may cause the death of the organism.BIOLOGICAL ORDER AND ENTROPY physical entropy is even more complicated. the structural negentropy of the system. Nucleic acid therefore contains the blueprint for the [93] . The biologist feels that the functional order It is is an essential part of functional order is the living system. The value of the negentropy obviously does not give a measure of the efficiency of the system. And happens. would point by an example. As is a result of an accident. is the same. even in if the mutation is negentropy has remained the same. the negentropy of fected cell will die. and meaningless it from as will a purely thermodynamical point of view. The living organism is not only an Low system proposed Brillouin has probability and value are not synonymous. however. noninfected cell. It has ceased to be nothing has changed: the content consequence of the introduction' of the genetic material a cell/virus system is greater than the negentropy of the normal. a generalized negentropy principle that would take into account the value of the organization or information. As a of a virus. the altered organism is now unable to function normally and to reproduce. But the inalive. clear. the information. If. that is. than for the physicist. lethal. that the term "information" has for the biologist a different sense Remarks on "Genetic Information" The synthesis of a given specific protein is under the control of nucleic acid. be seen next." This illustrate this is certainly a very pertinent suggestion. For the physicist. But the mutation being lethal.

in the letters is know provided the number of contain the same is. that negative entropy. privileged. for each gene controls the synthesis of one given specific protein. Physicists account or a that the value of a message is not taken into measure of the information. the genes. The concept information. of the action of ultraviolet light on [94] . in a sense. But the biologist knows different from every other. contains information that negative entropy. "genetic information" refers to a given actual structure or order of the hereditary material and not to the negative entropy of this structure. included in the biological concept of genetic The gene is as a machine. and our position therefore. In an organism. The biologist currently speaks of information for the synthesis of a given enzyme. is have the same information. were organic molecules. or organization. that is. provided they the number of same content. The called genetic information. and the idea of quality. is the specific value.BIOLOGICAL ORDER orderly assemblage of amino acids into proteins. for evolution must be life also taken into account. for the biologist. Entropy and Evolution When order is functional order is considered. of probability has disappeared. But there is something more. for very biologists say that at the beginning there These were formed as a result good reasons. This should apply to all genetic information. the organization represents information. A theorem by Einstein random assemblage of letters both contain the same information. the same. It is believed that originated some 3 X 10^ years ago and that the organisms existing today are the consequence of a long evolution. But. and this is most regrettable. the hereditary organization has every right to be called information. In a machine. Yet. For the physicist. or structural negative entropy. Nothing is. Both the physicist and the biologist should be well aware of this situation. which that each gene is bases. positive is known about the origin of life. it appears that the biological considerably greater than the figure calculated by consider- ing only the probability of the base sequence in the hereditary material. for the production with blueprint is a probability 1 of a given specific structure.

As a result. producing new to certainly very different and better organisms that outgrew the original prototype. despite the fact that and a it contains and reproduces structural negative entropv^ very remarkable functional order. The Source In his remarkable of Orderliness everything that is book What Is Life.BIOLOGICAL ORDER AND ENTROPY simple inorganic compounds. The living system considered on a world-^\•ide scale cannot contradict the second principle. which today constitute the livincr world. and an organic gel endowed with catalytic properties eventually appeared. has the over-all entropy decreased The system organism is plus food might be visualized as as system but certainly not an isolated system. has doubled expense of an increase Is of entropy S. The first organism in turn started performing experiments. The it present biological order includes the historical experience its of the organism acquired during includes evolution that it evolution. and as a result the entropy increased. have been performed. at the same time structural and functional. a large number of species were selected. A large number of experiments. the catalytic "metabolizing" gel gave rise something able to reproduce itself. The organic molecules then assembled on some substrate such as colloidal clay. x'Vnd it is because is so highly improbable. As a result of a large number of experiments. This was the first organism. They necessarily correspond to an increase of entropy. This increase should be taken into account in any calculation of the negative entropy of the organism. When an organism metabolizes. Schrodinger states that going on in nature means an increase of entropy [95] . heat of the surrounding world is excreted. or increased? a closed AiV smaller or larger than AS? In other A\ords. due to the production of two at the organisms out of one. due to the degradation of energy. Let us consider again the equation ( Organism )i structural + food — >-( organism )2 + waste + heat The negentropy N. from any current living being. The organization of the actually living systems has been acquired at the expense of an increase of entropy. Everything unfit has been eliminated.

or orderliness. the source of orderliness is reasoning has been endorsed by Brillouin. The the real fuel for the maintenance of a biological point of A is steam engine is combustion of coal is the source of I movement of the machine. which is death. according to Schrodinger. according to Schrodinger." conserved and never gets food does not really matter. know that the formula "the organism feeds on negative entropy" has been applied to the maintenance of the living system. or organization. kept working by the combustion of coal. The organism can remain alive. it is only for the negentropy which losses from ical it and which is needed to make up for the work done. produces positive entropy. maintains its its order by "sucking orderliness" from environment. Schrodinger himself has is. Because biological order. since energy is lost. The only source of biological order maintenance as well as reproduction of biological order both depend on the same process. biological order. terms of negentropy. The organism. pointed out very pertinently that the characteristic of life is the prois duction of order from order." that is. It has been known for a long time that only an organism can give rise to an organism. or simple degradation processes in the living system. metabolism. reproduction being the other. For the animal. who writes. says Schrodinger. Negative entropy is. the the sunlight. orderliness [96] . This type of sources of orderliness are more-or-less complex organic compounds. that is. for the plant. but negentropy is the important The statements concerning negative entropy. a measure of order. just like information.BIOLOGICAL ORDER and that the living being continuously increases its entropy. "something very positive. namely. it not the source of the structure. can be expressed in Now. only by continuously drawing negative entropy from its environment. in Energy contained matter. thus approaching the state of maximum entropy. which is correlative to an increase of entropy. This is certainly all right." even when it maintenance. as life should be examined from view and also from a physical point of view. such statements alive by sucking applies only to as "The organism remains from its environment. But maintenance of the organism — that maintenance of biological order — is only one aspect of life. "If a living it can get due to mechan- organism needs food. might be misleading.

When light meets atoms. it has to perform oxidoreductions. In the original chemical bond. But it is clear that what is important in the molecule of sugar is not the structural negentropv of its atoms. one part work. Academic Press. Science and Inj orfnatio7i Theory. source The same seems to be true for the energy of the chemical bond. if it The as the steam engine does. [97] . Unless the energy of a chemical bond of food is utilized. as positive as negative entropy be. does not handle concepts of grade or logarithms of The organism energy of physicists I light or of handles atoms or molecules and the chemical bonds. L. New York. What is important is the energy of the chemicar bonds. have consulted decided that Schrodinger's formula was it did not make A general agreement was reached only on one point. as a result of a chemical reaction. the probability that these atoms are arranged in a certain order. and the other is potential entropy. whereas others claimed that sense at If all./ degraded. REFERENCES Brillouin. The organism probabilities.BIOLOGICAL ORDER AND ENTROPY wants to work. But. according to some physicists I have consulted. Negentropy is a grade of energy. such as the energy of light or of a chemical bond. But energy of high grade. just It might burn sugar. (1956). The energy of light allows the plant to separate hydrogen ions from the molecule of water. work is produced. for example. cannot be subdivided into positive and negative is available for entropy. When. some of the perfectly acceptable. It separates and binds atoms and molecules. Orderliness is a probability. Work. has to burn something. one cannot speak of free energy and of its entropic component. is produced at the expense of the energy of the chemical bonds. heat is pro- duced and a part of the energy degraded. as its it does not seem sound to state that the plant that uses light of energy feeds on the negative entropy of light. The organism is unable to make use of energy abstracto. with a correlative degradation of energy. /". Nevertheless. work is produced and energy is organism. The organism takes in food and burns it. fed with pellets of negative entropy. might even a physicist would succumb.

pp. Quastler. 251262. The information content of a bacterial cell. New York. Linschitz. (1953). Prigogine. H. (1958). University of Illinois Press. (1957). So7ne Principles of Energetics in Biochemical Reactions. [98] . 111. In Essays on the Use of Infor?/iation Theory in Biology. (1944)'. Paris. Cambridge. Wiener. M. Cybernetics. Hermann. University Press. I. edited by H. Introductioji to Springfield. E. I. What Is Life? The Physical Aspect of the Living Cell. Thernwdyna?nics of Irreversible Processes.BIOLOGICAL ORDER Klotz. Charles C Thomas. N. Urbana. Schrodinger. Academic Press. (1955).

so as their synthesis. a sequence of a few We have learned that nuckic acid. is a complex molecular society in which macromolecules and groups of macromolecules are interacting. synthesis of essential metabolites.CONCLUSION We small molecules. anism has developed. the stable. The an elaborate system of a feedback mechfunctioning of the organism reflects the [99] . the one cannot be produced The organism. is controlled by the product of their activity. The functioning of each enzyme or group of enzymes. are which of in turn the catalysts for the are. or better by the needs of the organism. or the cell. But. have learned that the very basis of is biological order. nucleic acid is the varvintT structure responsible for mutations and evolution. considered on the time scale of the world. Stability and variability reside in the is function of the genetic material hfe. This dual one of the important aspects of Nucleic acid is the blueprint for the synthesis of specific proteins. Each group possesses its system. own regulatory As a result of evolution. Nucleic acids and enzymes from in a functional point sense that view. same structure. of specificity and diversity. complementary macromolecules ^^'ithout the other. considered on is the time scale of the organism. All the groups and all the various regulating systems interact. permanent structure responsible for the specificity and for the reproduction of specificity.

a molecular tendency toward autonomy. enzymes. extracted from the cell — any structure. as a human being. Life can only be the appanage of the organism alive. I have In a acting. These viral is functions may be submitted to specific viral repressors. structural The genes of the host cell The pathological order the normal order is. and at the same time as a dependent part of a community. or the apparently consequence of toward an escape from cellular control. Waves and complementary aspects of the atom. Structures and functions are the complementary aspects of the organism. has to visualize himself as a person. Only organisms are The upsurge viral. structural genes. The study viral of viruses has revealed the existence of sequential controlling the functions viral genetic material autonomous reproduction of the and the synthesis of viral proteins. The genetic material of a virus sometimes under the control of the regulating mechanisms of the host. * * complex organized system.BIOLOGICAL ORDER high degree of coupling between regulating genes. which all macromolecules are interdependent The living organism is an integrated system of macromolecular its kind. Any [100] . Separated from its context — that is. just as may undergo structural recombination and exhibit functional interchanges.T. are sometimes under the control of the virus. as part of a whole. During three weeks. Normal and pathological entities types of dual order. whether cellular.I. The story of the cell/virus system is the story of the quadruple interactions between two different dual. operator genes. as an independent unit. cell is and metabolism. is just an organic molecule. repressors. is and functional. I had the privilege and pleasure to be part of M. is of molecular diseases. either a nucleic acid or a protein. subviral. in units. the units are necessarily interone of us. Such structures and functions able to reproduce particles are the a thing as living substance or living matter does not as a exist. The normal a structural and functional system of order. whole.

And I should like to express my admiration for this institute. disease. [101] . M.I.I. some powerful and efficient repressor-operator system. is But so far as can see.T. is perfectly healthy. an integrated system of strucI As I am a physician. the de- velopment of which has been so beautifully conceived by Karl Taylor Compton. This probably because the problem of the induction and repression is solved by some elaborate feedback mechanism. just like a living being. which no macromolecule can resist. I \ hich I already is somehow and a as suspected. namely. am interested in have tried to detect some sign of I pathological process. that M.CONCLUSION discovered something tures and functions..T.

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nthesize a few thousand specific macromolecules. an independent systejti of reproduction. cell as The term cell is it is common now used traits have tranthat the for the metazoal well as for the microbe. the cells of an organism. macromolecules have to work in co-operation. One category. cells of a multicellular organism differ from the microbial cell which is an organism itself. In this way. macromolecules endowed with life a specific structure and a specific function. The and reproduction is the Both the cell of a multicellular organism and the microbe are the ultimate units of integration and reproduction. Reproduction. dependent. they many essential features that the scended the differences. cell. and Assimilation Life. The differentiated cells are organized into and the tissues into organs. corresponds to parts of an organism. as a controlled by the organism whole. is sometimes is listed among the discriminative features From a biochemical point of view.INTRODUCTION TO BIOLOGY and from the brain tissues cells. This can be accomplished only because of the presence and activity of specific templates that are parts of the living system.

prerequisite for assimilation. Y. M. (1911). A Symposiimi on the Cheinical Basis of Heredity. 269-3 10.Bioche?nistry. Reproduction includes assimilation. R. New The Cell . (1957). Englewood CliflFs. Antony van Leeuwenhoek a?7d His ''Little Animals. to synthesis. (1959). Protistenk. and the interplay of and functions. J. true to type and to genetic continuity.BIOLOGICAL ORDER The formula Metabolism is "the living being reproduces its its kind" expresses the fact that each organism produces a own specific macromolecules. York and London. What Is Life? The Physical Aspect of the Living Cell. Nature of Life. McElroy. (1948). A. is and reproduction. E. The University Press. Prentice-Hall. A. Szent-Gyorgyi. C. an organism can reproduce only if it is endowed with the power to assimilate.. W.. Stanier. New York. The Microbial World. (1944). and Adelberg. Physiology.. In the last analysis.. C. B. assimilation is certainly a discriminative feature of Assimilation lation. The Johns Hopkins Press. the notions of learned concerning life. organism. [6] . Morphology. Press. C. Sons & Danielsson. Schrodinger. In sense. Dobell. re- production. correlative to life. when something has been Anyhow." John Bale. (1932). eds. Doudoroff. It corresponds. growth. But the words reproduction and asshnilation will take their full significance only biological specificity. eds. E. (1957).. E. D. C. and Glass... London. The principles of protistology. J. Baltimore. and life is correlative to assimi- As a matter of fact. and Mirsky. Academic Dobell. and assimilation cannot be separated. Cambridge.. Inc. this life. 23. at the molecular the level of the organism. Academic Press. Arch. A. assimilation precludes specific structures level as well as at and includes metabolism. N. REFERENCES Brachet.

grows. This is the basis of evolution. etc. absorbs food. of one. Thus the growth and division of a bacterium is not the growth and division of its molecules. but a molecule cannot divide. same building blocks. nutrition. A molecule can be split into fragments. But we are not interested in the external appearance of things. metabolism. and divides. and can only be.Molecules might aggregate. which is two a microorganism. The specificity of an organism in the last analysis is. The statement [7] . This means that each individual always produces identical macromolethese species are constructed with the cules. What is reproduction at the molecular itself level? The But all living world is composed of thousands of species differing in their size. in the invisible essence of things. in We are not interested what we see but in what we do not see. . shape. some of which are inheritable. What is is it is biological specificity? How is the information stored which responsible for the biological specificity and diversity? How reproduced? Everybody knows that organisms undergo variations. A bv molecule is the smallest unit quantity of matter which can exist and retain all the properties of the original substance.THE PROBLEMS Organism and Molecules A bacterium. This process is metabolizes. but each fragment is necessarily different from the original structure. Neither can a molecule sfrow. the consequence of the specific structure of its macromolecules. bacteria are thus called reproduction' bv binary produced out fission. chemical composition.

Molecular freedom viral diseases are a catastrophe. in a sense fortunate. In harmony. that dangerously threatened. the constitutive. As a matter of fact. [8] . of its own As a result. about the way the organism as a whole decides what it has to do when faced with the genetic material. for otherwise life would be boring. Yet molecule sometimes decides to get rid of the shackles of co-ordination profit. In an ideal molecular society. all is sorts of molecular diseases are produced. When a molecule develops strong personality and refuses to submit herself to the common rule. Freedom is essential for the development and blooming of the human being. tion. How has the organism solved the problem of intermolecular communication? An organism rarely lives in a constant environment. is. are submitted to a dual action: they are controlled which is responsible for heredity and evolusame time they are subject to the action of the environment. of course. Each molecule Each molecule must be able to receive messages and must be disciplined enough to obey orders. its molecules. And one would like to know something about the interaction of hereditary and environmental factors. molecules have to work in know what the other molecules are doing. a what happens normally. When the environment varies.BIOLOGICAL ORDER that an organism reproduces true to type are is therefore in true only statistically. the organism has to cope with the variations. for the organism. Everyone knows This is that things are not always as they should be. dependent parts of the organan organism. molecular life. What the molecular basis of hereditary variation? An has to organism all is an integrated unit of structure and functions. and to work for diseases. the organism This is is each molecule works for the community. its experience. historical The structure of the organism includes past. But as part of an organized society the human being has to accept a certain number of restraints. I mean. Molecular a which only one aspect. by and at the a given situation. pose number of problems that we shall have to discuss. Heredity and is variation combined its the living system. ism. a This is true for molecules.

It was from the enzymes can be extracted only one chemical performs thus learned that a given enzyme to be catalyzed by reaction. and the word order has hardly been pronounced. The energy needed for the syntheses is provided bv^ the oxidation of food. and the energy obtained is stored as high-energy bonds. As everyone knows. is Biological order dual. In order to synthesize tryptophan. Syntheses take place. activity by themselves. A living being a dual system of order. We A have to know what this dual system is. each reaction requires energy. — COOH. Biological and it is especially a sequence in space a?2d time. synthesis of each building block is a stepwise process. Food is is seeded Chemical reactions are performed. Metabolism bacterium in. respectively. each enzyme. and finally each atom is exactly where it belongs. One atom is attached to another. sequence or succession in space or time. is taken in a proper nutrient medium. orders are is static and dy- namic. as a Order may also all that. for example sugar. carboxylated. a — CHo. for example in the form of adenosine triphosphoric one again mediated by a specific acid.THE PROBLEMS The Subject Our subject is biological order. Structural and functional the complementary aspects of the living being. The organism "burns" sugar. In a biosynthetic chain of reactions. Other enzymatic Some proteins have an help of a specific coenzyme. known concerning is the synthesis of or a intermediary metabolism. each reaction has a specific enzyme. [9] . As an The example. order is the fixed arrangement present be considered order is in the existing constitution of things. or aminated (attachment of. Most of enzymes work only with the the cell and purified. The oxidation of food and the storage of energy involve a series of chemical reactions. is Today. This the metabolism. molecule methylated. structural and functional. almost everything building blocks. a or an —NH2 in group). Enzymes are specific proteins. the biosynthesis of the amino acid tryptophan is depicted Figure 20.

what marvels there are in so small a creature." he wrote. "Dear God." The fact that life could not be something simple has been suspected for almost a century. In fact. the state of our knowledge. and size increases. The bacterial machine works. Claude Bernard produced an impressive statement. or perhaps the state of our ignorance. the bacterial cell. is life. they were cubes. a thousand packed into 1 milliliter. "by which one attempts to describe the protoplasm. Anyhow. cell the problem seems formidable and the situation hopeless.000 species of macromolecould be is one thousandth part of a millimeter. In his famed classical book. that is. And. itself divides. But for us. as the bacterium is devoid of brain.* corresponding to a volume of 10~i2 milliliter. not be concluded that knowledge and ignorance are synonymous. 1 /i A contains some 2.of them. machine of around 1 diameter. thus giving rise to two Simplicity or Complexity a well-known physical chemist interested in wrote this remarkable sentence. t If billions. manufacturing more of their far from simple. and finally the bacterium daughter cells. Les phhjomenes de la vie cormnuns aux animaux et aiix plantes {The Phenomena of Life Commoji to Afiimals a?id Plants). t a few thousand specific molecular species are at work. Claude Bernard certainly had no idea of how much more complicated protoplasm really is. synthesizes. however. And we would rather be inclined to say with Antony van Leeuwenhoek. is illusive. who in 1676 discovered the bacterial world. grows. lO^. the suffering human beings who try to A few years ago bacterial physiology penetrate the intimate nature of small." This was. [10] . In this specific kind. The nucleus divides. it has no problems. "The formula C18H9NO2. around 1875. Unity At * first sight.BIOLOGICAL ORDER As a result of the functioning of the enzymes. the protoplasm is much more complicated. the bacterium its synthesizes more enzymes. It should. "The structure of the bacterial cell is simple. despite being fx. and divides." This is true of course from the bacterial point of view.000 to 5.

Nucleic acids are made of four nucleic specificity is What will be learned later. the metabolism essentially the same in each Unity of coviposition: the main macromolecules of all living beings are composed of the same small molecules. nature has made use of a strictly limited number of building blocks. only necessary to For the time being. the problem of heredity. Uvity of pla?i: cules. in order to build the immense diversity of the living systems. each cell possesses a nucleus imbedded in protoplasm. know that the . gories of different organisms. one discovers unity. The problem of diversity of structures and functions. Yet.THE PROBLEMS Moreover. nature has produced an immense variety of catewhen the living world is considered at the cellular level. it is most important macromolecules are nucleic acids and proteins. and the prob- lem of diversification of species have been solved by the elegant use of a small number of building blocks organized into specific macromolecules. is Ufiity cell. of functiojT. For.

composed of twenty its An amino acid is.BIOLOGICAL ORDER bases. aminated acetic acid. for example.I \ C— C — C — COOH H N c H H NH2 V" . an aminated Glycine. In proteins./ " C^ I. as indicated is by name. thus forming nucleotide. thus forming long chain (Figure The amino acid. by phosphodiester bonds. analysis of proteins reveals that they are acids. A molecule of a phosphoric acid attached to each sugar. each bound to is a sugar to form a nucleoside. a The nucleotides are united 1). H H " \.

THE PROBLEMS molecule is for doing precisely endowed w ith a specific function. We may admire it. the living system did not perform its task. >/ [13] . The machine is built what it does. but we should If not lose our heads. We have simply to learn how it performs task. its it would not exist.

Schrodinger. in this fundamental book. tree is The a living reality. the chromosomal fiber. his well- known Esthetic. In its germ the determinants which was not already determinants exist as potentialities. the British philosopher Herbert Spencer. says the German philosopher. I have a suspicion that of the stu- somewhat unfamiliar with Hegel's Esthetic. Starting from small molecules — wrote Schrodinger in 1944 — it is possible to build large aggregates without the dull device of repetition. in his Principles of Biology. stated that in every organism the total of hereditary properties is determined by distinct physiological units which are formed by the assemblage of chemical units into immensely complex compounds. may be called an aperiodic [14] .THE HEREDITARY ORDER: GENETIC INFORMATION Birth of the ' Modern Concept Hegel published a small fraction In the year 1835. Whatever the case may be. even with the microscope. In 1864. and yet Nothing is in the germ one does in the tree not see anything. among a number of irrelevant remarks. every stone. in the germ. a crystal. The most essential part of a living being. the problem of dents here could be life is pertinently discussed. presenting the same solidity of structure as a crystal. plays an individual role not entirely equivalent to that of the others. Yet the most extraordinary biological intuition was that of a physicist. The atoms forming a molecule are united by forces of exactly the same nature as the numerous atoms which build a true solid. irrelevant for us today. as existing in the We can visualize the germ as extremely simple forces. In a complicated organic molecule. every group of atoms.

and not until 1953 that its it disclosed. The tions. how a molecule of hereditary material reproduced? Necessity of Variation The true. or ides. they were rediscovered inde- Hugo de Vries. is Each ide con- tains determinants. composed of biophores. which contains the chromosomes. according to They are relatively large Weismann. But Mendel's laws in remained ignored pendently by place on the material until. The chromosome. which means "colored bodies. and they stain heavily — hence their name. which all is composed of spherical bodies. in 1892. that to the unit of genetic material. Chromosomes have been known to cytologists for a long time. represent a different character. and each determinant is. Schrodinger adds that we believe the whole chromosomal fiber to be an aperiodic solid. And Weismann. that to multiply the specific biological order. would probably have disappeared long ago as the result of the drastic changes undergone [15] .THE HEREDITARY ORDER: GENETIC INFORMATION crystal. This was the beginning of formal genetics. and Tschermak. The biophore corresponds to the actual gene. But identified structure was was Each gene was assigned a specific was not until 1944 that the genetic chemically. is the cellular organelle which is responsible for the maintenance and transmission of the inheritable characters. Correns. understood that the inheritable characters were located in the chromosome. chromosome. Haeckel realized that the nucleus. and easy to stain. It is. the unit of life — has been defined as an independent system of integrated structures and funcis a highly complex system. able to reproduce its kind. If its kind is of course but only the primitive living being had life always given birth to an identical system. What is is the molecular basis of biological order and specificity? is How order stored in the is organism? If a molecule unable to divide. living being — the organism." As early as 1866. statement that the organism reproduces statistically true. The quantitative study of the transmission of hereditary characters in had been started 1865 by Gregor Mendel. 1900. the unit of biological order.

which a I would dislike to call final. which splits lactose into glucose and galactose. that is. quite obvious individuals belonging to one species were. Each time such situations it is found that the gene mutation. hereditary change. identical. or the hereditary change. are unable to oxidize amino acid by the normal route. a plant producing red flowers may modified offspring. some human mutants. When examined. Some human beings. responsible for the stepwise synthesis Human this beings normally oxidize the amino acid phenylalanine. phenyl- [16] . which is Mutations still take place nowadays. that if all It I suppose. It is admitted that the essential cause of evolutionary variation has sudden. later. as highly improbable systems. can only be the result of a stepwise process. found that one of the enzymes missing. Or a bac- terium synthesizing the amino acid tryptophan gives are analyzed. is acquired or is lost. evolution. place between the beginning. again. is. producing white flowers. wild type of another bacterium synthesizes the amino acid methionine. Living beings. the ability to synthesize a given enzyme. The biosynthesis is mediated by a series of enzymes. or mutants. the so-called a given bacterium is able to synthesize the enzyme ^-galactosidase. As a consequence. a given specific catalyst. there would be no such thing as genetics. corresponds to an alteration of an enzymatic system. Here the bacterium may produce mutants it is is unable to synthesize methionine. But wild type. Each enzyme corresponds to a specific gene. on the study of differences. rise to off- spring unable to synthesize tryptophan. or let us say an initial and a given. And the change is hereditary. As a result of a mutation. been the gene mutation. This the Beadle-Tatum law: one gene controls the synthesis of one enzyme. would be difficult to understand heredity. of it may The give rise to mutants devoid of this enzyme. Evolution is the sum of the alterations which is which have taken primitive state. The original. The principles of genetics are based differences.BIOLOGICAL ORDER by the primary environment. of transmissible give rise to For example. Let us give a few concrete examples. and would it is remain. In the absence of mutations. is transmitted to the offspring. So most fortunate that mutations do it exist. each responsible for a given step in the sequence of reactions. present state.

nature of the glucidic subunits are The polysaccharides are easy to identify. polysaccharideless bacterium. the Piieimwcoccus.Q. that to identify the nature of the entity controlling the production of an enzyme. unable to think. responsible for the the phenylpyruvic Long before the specific alteration responsible for a mutation had been chemically known. Material Identified Transformation. of these persons is is. The Genetic capsule. or which is a specific polysaccharide. They are idiots. This was accomplished by the study of bacteria. generally lower than 10. for a polysaccharide injected into an animal elicits the formation of a specific protein called an antibody (Figure saccharide combines specifically with the polyand not with polysaccharide II. the site of the biological alteration. Alany bacteria possess an outer layer. called ge7}es and described the of The problem was of course is. had been located on the chromosome. a Piieim703). and this information can be transferred to a bac- [17] . and long before the specific responsible enzyme had been identified. originally type II bacterium into a capsulated type III. The extract the of type III contains the "information" for the synthesis of type III polysaccharide. This however. originally of type in the for example. not important the in this analysis. Something present in extract of type III bacteria has trmisfoiined a capsuleless. The responsible as units chromosomal structures were variation. The agent of pneumonia. What is important is that enzyme present in normal human beings and normal pathway of phenylalanine is missing in idiots. produce its specific polysaccharide. is grown presence of extracts of type III bacteria. Sometimes. as already stated. The I. exhibits many varieties or types. It agglutinates bacteria of type III and not of type II. a mutation. The capsule of each of them is built of a specific polysaccharide in which the difl^erent. An anti-III-antibody III cocciis loses the ability to this capsuleless. to identify substrate of the hereditary variation. as a result of a sudden change.THE HEREDITARY ORDER: GENETIC INFORMATION pyruvic acid appears in the urine. When II. one recovers capsulated bacteria of type III.

BIOLOGICAL ORDER -^Antibodies agoinst type m ' ' .

endows it with endows it with a nucleic acid of given physiological potentiality. Desoxyribonucleic acid. Many bacteria have been transformed with DNA coming from a closely related form. multiplied and regularly transis But this specific nucleic acid not directly for the synthesis of the polysaccharide. a transforming principle can be extracted again. con- three types of molecules: (a) purine and pyrimidine bases. Avery. When a pure." This principle could not be metaphysical. The Finally. In all the cases of transformation studied so far. Thus the the specific is transforming principle. in a obtaining was Each fraction was tested for its transforming ability. the bacterium produces a specific a enzyme that can perform specific reaction. This is not a unique case but one particular example of a general phenomenon. it had to be an organic molecule. desoxyribose. MacLeod. or DNA. Once a gives rise bacterium has been "transformed" by a specific DNA. From the them. or DNA. [19] . They duplicate it in turn. and McCarthy succeeded in pure active substance that was identified as desoxyri- bonucleic acid. This information was called the "transforming principle. nucleic acid controlling synthesis of a specific substance. viruses can be separated The a from the protein components. that is. it to transformed offspring. Structure of the Genetic Material Desoxyribonucleic tains acid. in this case the synthesis of a specific poly- saccharide. the introduction of the desoxyribonucleic acid of the donor bacterium into the recipient organism the capacity to synthesize a given enzyme. specific viral nucleic acid penetrates specific viral proteins and specific viral infectious particles are produced. When it is present.THE HEREDITARY ORDER: GENETIC INFORMATION terium that was originally type II. and (c) phosphoric acid. naked. mitted from responsible cell to cell. bacterial extract containing the transforming principle fractionated. cell. The nucleic acid of viruses plays the same role in heredity as the nucleic acid of bacteria: both control the synthesis of specific proteins. 1944. (b) a pentose. The daughter bacteria have inherited the specific information. This conclusion has been con- firmed and extended by the study of viruses.

BIOLOGICAL ORDER

The two
idine
bases,

purine bases, adenine and guanine, and the

two pyrimFigure
4.
1

thymine and cytosine, are represented
3.

in

Each base

is

attached to a molecule of desoxyribose in position

carrying a phosphorus in position

This

is

a unit, a

desoxyribonu-

cleotide (Figure 1). In the nucleic acid, nucleotides are linked

by
1.)

phosphodiester bonds.

The

result

is

a

long chain. (See also Figure

adenine

thymine

guanine

cytosine
H

-H-K

'

H

\^~^cN
('sugars
C

\-/
C
//

\-H/

{ \

Vh
/

\
O'^UGAF

nh

Figure

4.

The Four Nucleic

Bases in the Desoxyribonucleic Acid.

is

Each pvrimidine base is bound to a purine base by hydrogen bonds. Adenine bound to thymine, guanine to cytosine.

Thanks

to

James Watson and Francis Crick, the structure of

DNA

has been disclosed.

DNA
is

is

a

double polynucleotide chain.

The two

by hydrogen bonds between the bases. bound to a purine base: adenine to thymine, guanine to cytosine. Thus in the double DNA helix, the ratio of adenine/thymine and of guanine/cytosine is equal to 1. The two
chains are held together

A

pvrimidine base

chains run in opposite directions.

They

are twined

around each

other (Figure 5).

[20]

THE HEREDITARY ORDER: GENETIC INFORMATION
This structure
is

now

firmly established.

The

evidence derives
(titration

from X-rav

studies as well as

from physical chemical data

curves, light scattering, viscosity, sedimentation, kinetics of break-

down by gamma
is

rays and enzymes).

Thus desoxyribonucleic acid is a double helix. Each helical chain made up of a chain of desoxyribose phosphate with one base
Ribonucleic
acid.

attached to each sugar.

DNA
is

is

the genetic material of
a

all

animals,
viruses

plants,

microorganisms, and of some viruses. But in

few

the genetic material

ribonucleic acid. Ribonucleic acid, or

RNA,
it is

has the same fundamental structure as
nucleotides.

DNA:

it is

is

a

long chain of

But whereas the sugar

in

DNA

desoxyribose,

ribose in

DNA
double

is,

RNA. Moreover, the pyrimidine base in RNA, replaced by uracil. Finally,

thymine present in whereas DNx\ is a

helix,

RNA
The
by

is

a single helix.

The two
information.

nucleic acids,

DNA

and

RNA,

both carry genetic
singularity

substrate of genetic information can only be a

feature shared
is

these

two

nucleic acids.

The common

obvioush" the linear arrangement of nucleic bases.

The chemical identification of the genetic material by Avery, MacLeod, and McCarthy has been the great discovery of modern biology. It was soon followed by another great discovery, that of the molecular structure of the desoxyribonucleic acid, the WatsonCrick double
virology,
helix. Both discoveries catalyzed the extraordinary development of genetics, cell physiology, cell biochemistry, and

which have now merged

into a

new

integrative discipline:

molecular biology.

Figure

5.

The Watson-Crick Model

of the

DNA

Double Helix.
are constituted

The two

chains run in opposite directions.

The backbones

by

desoxyribose molecules united by phosphodiester bonds. The nucleic bases of one chain are united to the bases of the other chain by hydrogen bonds.

[21]

BIOLOGICAL ORDER

The Molecular
Enzymes
species

Basis

of Enzymatic Specificity
are proteins. Proteins are
If,

of amino acids (Figure 6).
are organized

into

composed of twenty species two protein molecules, these twenty one and the same sequence, the two
in

Figure

6. The Molecule of Ribonuclease (from The Molecular Basis of Evohitio?i).

C. B. Anfinsen,

molecules are necessarily identical.
tein
is

The polypeptide

chain of a pro-

folded, but

it

will be admitted that folding

depends on the

sequence of amino acids and that two identical polypeptide chains

[22]

A given amino acid. and of course in the proportion of each of the twenty molecular species. except the terminal one. Fragment of a Polypeptide Chain. etc. at a specific peptide of the protein. Thus. is necessarily unique. The unit of function is the whole protein molecule.THE HEREDITARY ORDER: GENETIC INFORMATION same way.h CHg H HCH H CH2 H III CH2 CHg I I COO- CONH2 CH2 I \l + H3N^ I CH. differences in proteins in the sequence of amino acids which constitute the polypeptide chain. A long polypeptidic chain can be arbitrarily subdivided NH3 + OH ' CH2 C^ ' A 1 CH ' CH2 CHg I f-H 1^1 I c. When an enzyme acts. is necessarily associated to its left and to its right with another amino acid. the same peptides is may be repeated. being a specific molecule. Many molecules of the same amino acid may be present in a given protein. For the same groups of amino acids. the substrate attached at a specific site. the molecule. Their functional value depends on the sequence of all amino acids. In a protein. into peptides: di-. Each enzyme. tetra peptides. c K/ c CH2 H \c C II \ / c C II H \l ")c H \l c I c C ^C II — OH I N — OH I N —N OH I C II N I C II N I N II OH glycyl OH glutamyl OH III leucyl isoleucyl tyrosyl cysteinyl asparogine Figure 7. thus forming a tripeptide are necessarily folded in the can be due only to differences (Figure 7). no function can be assigned to any isolated amino acid or group of amino acids. But this specific peptide. This does not mean that in different species the same enzy- [23] . individual if isolated from the rest of would not be enzymatically active. tri-.

is antigenically (structurally) different in bacteria. If only one. synthesizes only a limited number of proteins. or group of closely related species. If just a were enough to take care of one amino acid. The enzyme /3-galactosidase. Now a protein is a specific sequence of amino acids. the The statea ment that each enzyme is unique applies only to the structure of given enzyme of a given species. in yeasts. a practically infinite array of specific proteins can be synthesized. is important is that proteins are synthesized on a nucleic acid plate. for example. and that is too low. Protein Synthesis: The Code DNA. with twenty different species of amino acids present in various numbers and proportions. One problem. with the is exception of some viruses. however. thus contained in has first seems probable that DNA to manufacture an RNA What tem- template and that proteins are synthesized on this template. A given organism. step by step. An organism is apparently unable to manufacture incomplete proteins. of the amino acids entering into the constitution of a protein is missing. and in animals. hydrolysis of the ^-galactosidic link. How does a sequence of four nucleic bases code a sequence of twenty amino acids? The problem of the code has been often discussed in recent years. If we assume that the process does not [24] . Things happen as if the synthesis were a sort of zipper-like process. It should be added that the synthesis of a protein is an all-or-none any one. starting at one extremity of the template and continuing process. The tem- plate therefore has to be specific. the number of permutations would be 64. the number of permutations would be 16. no protein synthesis is possible.BIOLOGICAL ORDER matic function cannot be performed by different proteins. We shall closely follow Francis Crick's excellent review. Let us pair of bases consider a sequence of nucleic bases: ACBDACDBCAD start at the C C B A C. has many solutions. It The highly hereditary information for the synthesis of proteins. Thus. If three bases were required and if any set of three bases could take care of one amino acid.

A A given amino acid can be coded only on one of "good triplets" being twenty. ACB.. the difficulties are the same. valid. BCA.THE HEREDITARY ORDER: GENETIC INFORMATION end. the total number This means that 44 triplets out of 64 do not make sense. the possibility^ that a to a is given sequence of amino acids. on one of the chains rectly built only. given triplet corresponds to only one amino acid. Moreover. If one is composed exclusively of "good" triplets. A given sequence of bases necessarily gives this hypothesis. ABA. ho\. and the end results will be determined by the point of departure. The good 8. Crick's Tentative Solution of the Coding Problem. is Any other does not others are bad triplets. Thus no overlapping rise possible. ABB. In single chain. according to the discussed code. etc. Crick. But the double DNA helix is from two complementary chains. No bining the adjacent bases of is good triplets can be obtained by comtwo good triplets. — which make sense — are represented in Figure All the A B C D Figure 8. the process can start in A or in C or in B. If there is a possible overlapping. Griffith. template a The protein may be synthesized di- on DNA is not excluded. Any make combination sense. and Orgel have worked out 1. the other chain would necessarily have a large fraction of "bad" triplets and would therefore not make sense.do the amino acids is know which template to read? If there no overlapping. a code possessing the following properties: triplet. there are 64 permutations. it would therefore be. . If proteins were synthesized on a DNA template. triplets 2.

even more refined data were oba bac- tained. has been learned that the nucleic acid is the substrate acid. they mate. however. Matthaei. Whether a sequence of three or four nucleotides is involved is not yet known. the coding problem considered open. Academic Press. ability of Two closely linked structures have a high probIt remaining joined. The spontaneous mutation rate in a bacterium or in * Since the Compton Lectures were delivered (March. 15881602]. pro"messenger. It is clear. By the use of mutagenic agents. is Anyhow. Moreover. work- ing with bacteriophage. 1961. DNA RNA [26] ." /. becomes attached onto a nonspecific cytoplasmic ribosome. New York. Sci." This messenger. template a. Thus: (a) the genetic information is contained (b) the code is a sequence of nucleic bases. that the whole code will soon be deciphered. Nirenberg and J. It has been learned that the structural gene. The code is now being deciphered: a synthetic poly-uridylic acid contains the information for the synthesis of a protein having the characteristics of poly-L-phenylalanine. our knowledge concerning the synthesis of proteins in general and the code in par- ticular has increased considerably. 1960). b. 47. 3. Proc. that the unit of variation was of the order of three to five nucleotides. to put in order). MoL Biol. of mutations. It is the system ribosome + messenger which assembles the amino acids into a given sequence: diataxy (etymologically. Acad. 318-356.* Mutation The nucleic it in the nucleic acid. detached from the chroduces a specific mosome. The longer the distance.6 to 2. is. A synthetic poly-cytydilic acid takes care of L-proline [M. the greater the probability of recombination. See Jacob and Alonod: "Genetic regulatory mechanisms in the synthesis of proteins.BIOLOGICAL ORDER Finally. the specific carrying the genetic information for the synthesis of a given protein. W. and Jacob and Wollman: Sexuality and the Genetics of Bacteria. thus producing a specific protein. When two same cytoplasm. It happens that the probability of recombinations is related to the distance between the two difl^erent chromosomes multiply in the factors. (1961). was found by Seymour Benzer. H. and recombinants are produced.8. How this is re- flected in the constitution of enzymes is not yet known. as entirely for the time being.'Nat. that of hereditary variation. The smallest unit of gene variation can be measured by recombination experiments. varies the ratio (adenine + thymine)/(guanine + cytosine) with the bacterial species from 0. (1961).

THE HEREDITARY ORDER: GENETIC INFORMATION relatively low: one ov^er one hundred thousand is (10~5) to one over one hundred million (10~8) per generation. Among these "mutagenic agents" are X rays.-. or naked viral nucleic acid are exposed to process nitrous acid. cytosine pairs with guanine.CH3 ^C_N > N^ J:c /O C H-N /C C /CH3 '. such as nitrogen mustard. This means that the deamination of one pair. an adenine/thymine of deamination._/ \ \ / ^N r q/ C \_N —H ( hypoxanthine deam. A4any agents are known to increase the probability of mutation. The is kinetics of the a one-hit process. base enough to produce one mutation. or animal viruses. which is thus converted into hypoxanthine. or bacteriophage. bacteria.ne) thymine CH3 H N deamination \ / H \ hypoxanthine H H cytosine guanine H cytosine \ C— N ^N^. noted oden. The original base pair adenine /thymine has a result been replaced bv the base pair guanine/cytosine as of the action of the nitrous acid. At the second rephcation. Nitrous is. >— . mutants are produced. for example. and teriophage nitrous acid. deaminates nucleic bases. ultraviolet rays. and various chemicals. Let us consider. H Odenme thymine N ' >-< < }-< "-\_/"^" >-% /-" \ /"' \/^-\ . organic peroxides. adenine 9). as will be seen later. when is acting on nucleic acid. that takes off their NHo group. When. or plant viruses. is As a result transformed into hypoxanthine (Figure The replication of the DNA double helix. [27] .0 H —N / C / C N— H Z ^N >—< N \—N . The Mutagenic Action of Nitrous Acid (from Robert Lavalle). reveals that induction of mutations acid. At the first replication. for example.Cy <.CH ^c )^-" \ C/ /" "^ \-H after one replication and tautomerization after a second replication Figure 9. Nitrous acid deaminates adenine. hypoxanthine binds cytosine.

" The "mutated" protein? protein. to a given wild-type gene and to this gene once has undergone mutation? Hemoglobin. nothing more. of a complementary strand. the original base pair. By But the differences have been traced back to one amino acid. Amino Acids Sequence Hemoglobins. the hemoglobin can be cut into small peptides. which in turn will bind guanine. In a hereditary blood disease of man. the Table I. the protein of the red blood of oxygen. one of these peptides is different in the normal and mutated proteins. Thus. What the result of a gene mutation in the What a is the difference between the proteins corresponding. the pair guanine/ is one of the daughter cytosine. hence the sickle name of different the disease: sickle-cell anemia. replaced by in fact a so-called "point mutation. will bind cytosine. hypoxanthine. is in helices.BIOLOGICAL ORDER essentially the formation. As discovered by Vernon Ingram. mild digestion. in Normal and Abnormal Peptide fragments of normal hemoglobin (A). The normal human red cell is a disc: it looks round when it lies on its flat surface. associated with the so-called "hemoglobin C disease. The in a hemoglobin is from the normal hemoglobin number of properties (Table I). has been the subject cells responsible for the transfer of extensive studies. The new base. and hemoglobin (C). We is have learned that nucleic acid con- trols protein synthesis. In the sickle hemoglobin S. A gene mutation." Histidine . by each of the strands. the red cells are reduced in number and exhibit the form of a sickle. as a consequence of a deis amination by nitrous acid. it respectively. adenine/thymine. sickle-cell hemoglobin (S).

THE HEREDITARY ORDER: GENETIC INFORMATION
glutamic acid
acid
is

is

replaced by valine. In hemoglobin C, glutamic
lysine.

replaced

by

Thus we have

learned that nucleic acid

is

the substrate of heredi-

tary properties, and that the genes control the synthesis of specific
proteins, or of specific enzymes.

We
is

know

that the sequence of

nucleic bases in the genetic material

responsible for the sequence

of amino acids in the proteins.
base pair

We know that the

replacement of one

by another may be
difference

responsible for a mutation.

We

also

between the normal and the mutated protein may be due to the replacement of one amino acid by another. As a consequence of the replacement of one base pair by another in the gene, one amino acid is replaced by another in the protein. The sequence is changed, and as a consequence the folding of the polypeptide chain might be changed too. The nucleic acid,
that the

know

the genetic material,

is is

the substrate of the structural, hereditary
the agent through

change.

The
is

protein

which the hereditary

change

expressed.

Reproduction of the Genetic Material

DNA. The
that
is,

genetic material, or the genetic information,

is,

by
cell,

definition, transmitted

from the mother

cell to the

daughter

multiplied.

How

does one molecule of nucleic acid produce

two
of

identical molecules?

How

has nucleic acid solved the problem

acid is a double and thymine on the one hand and guanine and cytosine on the other hand are united face to face by hydrogen bonds. The double helix is thus composed of two complementary strands. Watson and Crick in 1953 put forward the hypothesis
helix in M'hich adenine

molecular reproduction?

Desoxyribonucleic

that each strand acts as a template for the organization of a

comple-

mentary one.

As

the

tion of the
in fact,

two strands of the duplex are complementary, the separatwo strands cannot be described as a binary fission; it is, an inequational cleavage, the separation of two different

But as these parts are complementary, each one, when producing its complement, reproduces the original molecule. The unit,
parts.

[29]

BIOLOGICAL ORDER
which is the molecule, is made up of two firmly bound complementary parts. The molecule of DNA cannot grow and does not grow. But by making use of complementarity it has solved the problem of molecular reproduction. By the template mechanism, a single strand of nucleic acid can produce a molecule of protein. It can also produce a complementary nucleic strand, which in turn can reproduce the original one. But only a molecule made of two complementary structures can reproduce directly its own structure. This is all right on paper, but how are things in the living system? An enzyme has been isolated from a bacterium that polymerizes desoxyribonucleotides. If these building blocks of DNA are added in a test tube together with the enzyme, is synthesized. The necessary condition, however, is the presence of a DNA primer.

DNA

The
In

original

DNA,
as

the primer,

may

be multiplied

i?i

vitro

bv

a

factor of 10.

DNA,

already mentioned, adenine

is

bound

to thymine,

guanine to cytosine.
guanine/cytosine, (adenine
is

The
equal

ratio

adenine/thymine,
1.

like

the
the

ratio
ratio

to

But,

in

bacteria,

+

thymine) /(guanine

+

cytosine) varies from 0.6 to 2.8

according to the species.

What
primers?
First:

happens

when

these various types of

DNA

are taken as

Second:
missing.

The ratio adenine/thymine and guanine/cytosine is always The synthesis does not take place if one of the bases

1.

is

Third: Each base can be replaced by an analogue, provided its hydrogen-bonding capacities are in agreement with the Watson-

Crick model.
Fourth:

The

ratio

(A

-f-

T)/(G + C)

in the synthesized

DNA

is

the same as in the primer. If the "nonnatural" synthetic primer does

not contain guanine or cytosine, these bases are absent in the newly

formed
Fifth:

DNA, despite being present in the test tube. When the DNA primer is heated, the two strands
is

separate,

and the rate of synthesis
Everything happens

markedly increased.
if

as

one of the strands would act

as

a

[30]

THE HEREDITARY ORDER: GENETIC INFORMATION
template. Experimental data obtained so far are in full agreement

with the Watson-Crick hypothesis.

more? Three mechanisms could be envisaged for the replication of the DNA. These are called conservative, semiconservative, and dispersive (Figure 10).
learn something

Can we

1 a

U

form another pair of hydrogen bonds with a third base. an enzyme has been discovered which binds ribonucleotides and organizes them in long chains. Its DNA was labeled with radioactive phosphorus. The formation of a complementary strand by the template mechanism would end in two chains with qualitatively different genetic information. an experiment consists of labeling the is needed. As the two chains of the double DNA helix are complementary. and is more generally not yet solved. the synthesis of proteins. The same type of problem is posed by RNA. It DNA of the parent organism and following the distribution of the label in the offspring.BIOLOGICAL ORDER In order to answer our question. Reproduction of RNA. A number of experiments perform. One could of course be active. 3. Only one of the strands of the DNA duplex produces an RNA the chain. the C of the sugar and the N of the base is to be always in the same 2. Bacteriophage was selected as the material. But it could well be that [32] . The conclusion was that DNA reproduces by the semiconservative mechanism. position. according to Stent. Each of the two strands of the DNA duplex forms an RNA chain. specific sequence of the RNA bases Each of the bases of the base pair of the duplex could. in agreement with the original Watson-Crick hypotheses. a given pair of the duplex bases would combine with only one base of the future RNA. The single-stranded RNA The is formed within the deep groove of the is DNA double helix. the two RNA chains would be different. RNA is single-stranded. Three mecha- nisms have been considered. Production of an RNA template froiM DNA. If the bond between pairs of the governed by the base DNA duplex. The first experiment by Cyrus Levinthal in 1956. 1. which is the genetic material of a number of animal viruses and of plant viruses. The problem of the mechanism by which DNA produces the RNA template for an RNA molecule.ed since on various materials seem to agree with this of this kind was done conclusion. and the concentration of hot phosphorus was measured in the daughter bacteriophage. the other being inactive. that is. Here again. the other inactive.

. Anfinsen." The key problem now REFERENCES The Molecular Basis of Evolution. B. The Macmillan Com- New York. Cambridge. (1951). forget the question of the replication of the ribonucleic acid and of viruses containing ribonucleic acid.THE HEREDITARY ORDER: GENETIC INFORMATION RNA is A — possesses the right sequence of triplets coding of amino acids and the synthesis of proteins. or that the single-stranded produces a double-stranded DNA. E. The mechanism would be the reonly one — let us say suitable for the RNA RNA verse of the synthesis of RNA from us." wrote Max Delbriick in 1949. New Dunn. Genetics in the 20th Century. (1947). desoxyribonucleic acid. one may perhaps suspect that the key problem of biology. how living matter manages is and perpetrate experience. Dyna77nc Aspects of Bioche772istry. physicist's point of view. "that the essence of life is the accumulation of experience through the generations. York. Anyhow. Conclusions The conclusion has been reached that each organism contains a structure. the problem not solved — not yet solved. Baldwin. C. then. John Wiley & Sons. A change "If it in the base sequence results change in the nucleic acid specificity. L. This structure contains the information for the pro- duction of specific templates and of specific proteins. [33] . pany. C. is from the to record solved. be true. which is the very base of heredi- tary order. DNA. The University Press. The specificity of nucleic acid is is the sequence of the nucleic bases. The other — let us say B — would be able to produce only a complementary strand A. Let for the time being. its reproduction is the production by each of the strands of a complementary one. and the sequence of the in a the specificity of proteins amino acids. ed. Because is DNA composed of two complementary strands. It could also be that a temporary double helix is formed. (1959).

Baltimore. eds. John Wiley & Sons.BIOLOGICAL ORDER Fruton. (1955). D. S. (1959). York. 17-26. Rich. [34] . The Johns Hopkins Press. Wagner. McElroy. John Wiley & Levinthal.. (1956). 42. Genetics and Metabolism. K. Nat. A. C. New York. and Simmonds. The mechanism of DNA repHcation and genetic recom394-404. Brookhaven Symposia in Biology. bination in phage.. General Biochejnistry. R. Proc. Sons.. Set. Acad.. S. and Mitchell. (1957). 12. Polynucleotide interactions and the nucleic acids. B. J. H.. P. New W. A Sy7t7posium on the Chemical Basis of Heredity. and Glass. (1953).

A bacterium contains about 2.000 genes and 2. the building blocks are synthesized step — the nucleic bases and the amino acids specific proteins — by step. are interdependent subunits. the double mother divides into two daughter cells. This DNA is reproduction. exist If the factory manufactures cars. which. that is. a given balance must etc. the building blocks are assembled in an orderly way and specific nucleic acids. In the first phase. or RNA. or something. doors. cars. wheels. When the genetic material has duplicated. they are [35] . acids Each enzyme is a specific sequence of amino which performs only one chemical reaction. has to co-ordinate and to orient the activity of the individual units.THE FUNCTIONAL ORDER The Problem Life has been equated with a complex hereditary order invoh'ing a specific sequence of nucleic bases. For the first process. are parts of a whole. Growth and multiplication depend on the completion of two sets of processes (Figure 1 1 ). The second process involves specific templates. only enzymes are required. This is growth. that depends on extrinsic factors. as parts of a whole. Thus more enzymes and more nucleic acids are produced. The units is.000 enzymes. contains the information for the synthesis of enzymes. the genetic material. The fiber of nucleic acid. between the number of motors. some- body. In the second into phase. the workers. Aioreover. brakes. In a factory involving the activity of a few thousand workers. the rate of production has to be adjusted to the con- sumption of it of the factory.

the essential metabolites (amino acids and nucleic bases). It has to synthesize all the in the right proportions in excess and in the right amounts. are synthesized. And waste decreases the chances aminoacids metabolism reproduction Figure 11. and Reproduction. The duced cell is a factory. The statement that the organism reproduces true type is [36] . A substance proin means waste. Metabolism. An imbalance may lead to cellular disease and even to live in a death. Moreover. Nucleic bases are organized into nucleic acids. especially between nucleic acids and proteins.BIOLOGICAL ORDER interdependent units. Food is metabolized. organisms do not constant environment. the Struggle for life. In nature. This is true especially for bacteria. The small building blocks. they encounter a wide variety of conditions. these poor beasts to are submitted to terrifying ill-treatment. different types of devices conbuilding blocks trolling order are interacting at various levels. The duplication of the desoxyribonucleic acid (DNA) is is the very basis of reproduction. prerequisite for protein The manufacture synthesis. A balance also has to be maintained between the various macromolecules. Amino acids will of ribonucleic acid the be patternized into specific protein on the ribonucleic acid templates. Growth. In a factory. And in the laboratory.

the bacteria are unable to grow in a medium containing carbon and energy source. galactosidase is synthesized only when lactose or a few other ^S-galactosides are present. considered: inhibition of As an introduction. The potentialities of the organism are under the control of the subject to the action of the environment. the integrated system of specific interdependent macromolecules. If lactose is supplied. The enzyme that controls its metabolism splits the /?-galactosidic bond and is therefore called /3-galactosidase. The induced synthesis of the enzyme lactose as the sole [37] . quantitatively as well as qualitatively. As a result of its is activity. When no lactose is present. two models will be one of induction of enzyme synthesis. but the functioning of the genetic material and of the whole organism is The organism as to has to cope with the environmental changes as well its ensure the harmonious interplay of in the enzymes. tose. Bacteria grown in a medium containing Bacteria in a synthetic grown glycerol are completely devoid of enzyme. Lactose induces the synthesis of galactosidase. the synthesis of the galactosidase starts after a lag of a few minutes. We have to dis- is informed of cytoplasm and in the outer world. however. Thus. genetic material. This a prerequisite for the utilization medium where the only organic compound is lactose contain the enzyme galactosidase — about 6.THE FUNCTIONAL ORDER Statistically correct. Lactose is a which glucose and galactose are united by a ^-galactosidic link.000 molecules per bacterium. How are the orders transmitted from one macromolecule to the other? What is the basis of molecular interaction and balance? The cell. In the absence of galactosidase. disaccharide Models of Enzyme Synthesis of the bacterium Escherichia coli are Some in strains able to utilize lactose as a carbon or energy source. and within an hour or so each bacterium has manufactured its full load of enzyme. lactose is hydrolyzed into glucose and galacof lactose. Two Induction. the enzyme is useless. the other of enzyme synthesis. will be close the mechanism by which the genetic material what happens studied as a functional unit. The enzymatic equipment reproduced true to type of a bacterium is is the far from being constant. But what It varies is genetic material.

Of course they have medium. building blocks. that is. When growing in a synthetic methionine synthase. What happens when the growth in the absence of methionine. synthetic The bacterium Escherichia coli is able to grow in medium devoid of any amino acid. the bacteria contain to. they are unable to synthesize methionine. [38] . or essential me- among them. to © cysteic C C II SO. Such a bacterium all its therefore able to synthesize tabolites. why this It is process has been. C ^ ^ _ COOH COOH COOH C NHj C — NHj COOH C I S — CH3 c I C I — NH. for a long time. and. COOH homocystine methionine METHIONINE SYNTHASE Figure 12. called enzymatic adaptation. And they are unable to synthesize grow. The enzyme methionine synthase is actually present (Figure 12). a is Repression. proteins. The Synthesis of the Amino Acid Methionine. Let us now consider the reverse process. now described as an induced biosynthesis of enzyme. the amino acid methionine. is The last step in the synthesis of methionine catalyzed by an enzyme called methionine synthase. For in the absence of methionine synthase. NHg acid—H* cysteine " C "t" ^ homosenne C SH NH2 C S C— OH .BIOLOGICAL ORDER increases the fitness It is between the bacterium and its environment.

sponsible as a result is of this excess.THE FUNCTIONAL ORDER medium is is utilized. The cytoplasm is limited by a thin membrane. we observe an adjustment of the enzymatic The cell tends toward a physiological balance. the bacteria are assayed enzyme has disappeared. But. A balanced state to an equal production is reached which corresponds and consumption of methionine. is surrounded by a cell wall whose function is mainly counteract the internal The bacterium to osmotic pressure. The development of our knowledge concerning the functional order has been possible. after for methionine synthase: the few hours. is Suppose now that the bacterium it methionine. Let us consider next the problem of the interactions of genes and of enzymes. a decreased rate of is. Whether we deal utilization of the carbon and energy source. the inhibition of called repression. This excess can be due either to an excess of enzyme or to utilization of methionine. And. bacterium and its environment. The result is the same. namely the amino acid methionine. thanks to the existence of nonhereditary. Our problem is to learn what the responsible mechanism is. system. a is supplemented with methionine? A labeled methionine methionine with a hot sulfur atom. Among the remarkable properties of [39] . Repression of enzyme synthesis thus increases the the fitness between with the lactose. The labeled methia onine taken up. and that is growing in the absence of produces an excess of methionine. The Genetic Control of Bacterial Metabolism Metabolism of lactose. the synthesis of the re- enzyme repressed. Escherichia coli will be taken as a tool. namely the sugar or with the synthesis of an essential building block. such as the induced synthesis of enzymes or the repression of en- zyme synthesis. The development of genetics and the chemical identification of the genetic material have been possible because of the existence of hereditary variation called mutations. not transmissible variation. that an excess of methionine. This means that in the presence of extrinsic methionine. This phenomenon. and as a consequence the synthesis of methionine decreases. the synthesis of the enzyme is blocked. enzyme synthesis.

" [40] . The two structural genes y and z control the synthesis of the /3-galactoside permease and the ^-galactosidase. bacterium. another for the sugar glucose. Lactose the by the model bacterium possesses Both enzymes are synthesized is an inducer. etc. o D — I ' permease cr •^gal I • I 1+ z+ y+ ^ > O o v^ • O O • O ^ /3. Our both permease and /?-galactosidase.galactosidase Figure 13. as in logic. the i /3-galactosidase splits lactose into glucose and galactose. another for the amino acid methionine. enzyme called permease.BIOLOGICAL ORDER the bacterium is selective permeability. Its is controlled by a specific per- role is just to take the molecules of lactose in the environ- ment and to transfer is them it is across the membrane. Accordingly. split Once lactose has been pumped into the cytoplasm. a Schematic representation of respectively. into glucose and galactose original enzyme ^-galactosidase. The permease /3-galactosides is responsible for the specific transfer of lactose and other the across membrane. only when lactose is present. The Utilization of Lactose. "based upon or using induction. A bacterium is not permeinside able to everything. lactose -^glucose + galactose —n . (Figure 13). The gene controls inductivity versus constitutivity. enzymes are said to be inductive. Inductive means here. the bacterium of And it was discovered that the penetration many small molecules is mediated by a There is given specific one specific permease for the amino acid tryptophan. a reaction in which an acetylation perhaps involved. the metabolism of lactose mease. Thus. another one for lactose.

to rise to hereditary vari- mutants belonging to various categories: bacteria able to synthesize /5-galactosidase (a) The may give rise to variants unable to perform the synthesis. (f o . This original model can give ants. the mutation is z+ ±^ z~.THE FUNCTIONAL ORDER Variations. z+ being the symbol for the character carrying the information for the galactosidase.

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^ = inductive I i+ I z+ y+ K I i i+ O = permease • = y5-galactosidase /5 o 1 O • 6 • • 6 • =constitutive . where i+ stands for inductivity. the mutation + i~. i+ in the absence of an inducer. being the symbol for the charis acter controlling inductivity versus constitutivity.BIOLOGICAL ORDER (c) The inductive bacteria may give rise to mutants that pro- duce /?-galactosidase and permease They i are called constitutive. i~ for constitutivity (Figure 15).

This gives a measure of the relative dis- tance of the genes. pair The normal gene ment of one base A point mutation. z+ . template. and i. b' I Figure 16. as a unit. mechanism. for example in a (a+ > a~) or in b (b+ > b~). As already Are seen. controlling the utilization of lactose. or corresponding of zipper-like 16). one finds: that the characters of the male can be transferred to the female.THE FUNCTIONAL ORDER The gene jugate. possesses the structure a+ b+. is the friction forces separate the partners. namely. 2. the determination of the frequency of exchange of one piece of a chromosome for another. Gene Mutation. When 1. and possible types of combination are found. as a whole. If this done at intervals. y+ . can mutate independently. [43] . a— b+ 1 mutation A „ mutation 8 . Let us supposed that a gene. When males are mixed with females. that the three "genes. the process lasting around one hour. can take place on various gene." z. and is conjugation interrupted. one finds that each gene of the male enters the female at a given fixed time. and that the various genes always enter in the same order. . This order can be verified by the classical tests of genetics. a replacesectors of the by another. these types of experiments are performed. these genes parts of the same functional unit. produces a protein. This allows us to locate the place of each gene on the chromosome. by a sort now consider the gene z (Figure The a+ b+ 1 normal gene . Our bacterium exhibits sexual processes. i+. or do thev reprethat the units sent different functional units? are different? How can we know its What justifies It is the use of the term "unit"? Altered enzymes. the all different genes. If the couples are put in a Waring Blendor. are closely associated or linked. they conof the male is The chromosome injected slowly into the female. sexual digression. y.

but instead a new. altered protein. to enzyme. Now the mutants a~ and b~ are different. the normal structure. The Activity of Normal and Mutated Genes. that is. that is. is present. It may have a slight enzymatic activity. that a fraction is. is present. one to the a+b+ corresponds for the the information synthesis of the Table II. In [44] . When the mutated gene z~. despite the presence of all the information. the enzyme is synthesized (Figure 17). The mutated gene z~ does not produce the original enzyme. Let us then mix the various z~ mutants. they can saturate (20 to 60 per 100) of the antibody.BIOLOGICAL ORDER structure "normal" gene z+. replaced by Some only fold. All this shows that the mutations affecting the gene z+ are mani- The proteins produced /3-galactosidase by the various z~ mutants differ from the enzyme and differ also in each of the mutants. As a result. others give incomplete cross-reactions. and any nucleic base. ture /?-galactosidase. Such a heterozygote is unable to manufac. can be replaced by another (Table II). The normal gene z+ produces the enzyme. Some z~ mutants give a complete cross-reaction. they are able to displace entirely the enzyme from the enzyme-antibody complex. whether a~ or b~. and in the protein one amino another. The male carries a gene a+b~ and the female a gene a~b + The zygote has two different sets of genes and is therefore called heterozygote for gene z. or be totally devoid of activity. An important fraction of the negative mutants z~ nevertheless contains a protein able to cross-react with the anti-/3-galactosidase antibody and to displace ^-galactosidase when combined with the antibody. whether in position a or b. no enzyme is synthesized. — normal enzyme (less "Normal" gene Mutation Mutation II I > active protein > active protein — modified enzyme — no enzymatic potent) > inactive protein activity sequence of bases acid is is replaced by another. So when the gene z+(a+b+). A gene is a sequence of nucleic bases.

THE FUNCTIONAL ORDER < d .

The male injects its chromosome female. Why that an inducer sometimes needed for the to gene to express tutive itself. When the bacterium necessary. it is One to takes a z+i+ female that possesses the right information for the synthesis of the enzyme. Another hypothesis has to be considered: the gene i+ manufactures a repressor.BIOLOGICAL ORDER Induction and Repression Generalized induction hypothesis. is i+ (inductive). is is is. then the zygote should synthesize the enzyme in the absence of an extrinsic inducer. and the female is z~i~ instead of the male. the enzyme synthesized whether or not an extrinsic inducer is present. the gene i~ being unable to Let us then admit that the gene i+ does synthesize a repressor. The repressor. the heterozygote z+i+/z~i~ starts producing /3-galactosidase at full speed. The reverse experiment is then performed. One takes a it z^i" male (Figure It cannot synthesize enzyme it because does not possess the right information. but. The into the male is z+i+ instead of the female. to inductivity. that is. if the function of the gene i~ is to manufacture an inducer. In the absence of an inducer. And the constitutive female cannot manufacture the enzyme because it is z~ . A [46] . an extrinsic inducer is. do so. unable males manufacture enzyme in the absence of an inducer. by definition. But possesses the constitutive gene i~. It tested. the female cytoplasm corresponds to the gene i+. could produce enzyme in the absence of an inducer if it were z+. manufactures an inducer This hypothesis can be easily 18). The gene z+ contains all of the information for the synthesis of a y8-galactosidase. Now and females are mixed. Male and as the female are again mixed in the absence of an inducer. Things are more complicated than was thought. make use of its potentiality is by manuit facturing the enzyme? The simplest hypothesis that the consti- bacterium does not need an extrinsic inducer because itself. When the bacterium is i"~ (constitutive). If the theory is right. It does not. In the zygote type I. the male cannot facture the manu- enzyme because it is inductive. But the bacterium that possesses the gene z+ does not necessarily manufacture the enzyme. being i+ (inductive). As soon z+i+ gene from the male has penetrated in the z"~i~ female.

THE FUNCTIONAL ORDER X X X X X ^ X X 2+ X X 5^ X X X X ^X X "" X ^ X X X > X X v- X I I cr zygote I .

should be responsible for the manufacture of a repressor. which. in is the absence of an inducer. This If. and the enzyme synthesized. is No repressor is present. But the male has introduced at z+ introduced the same time the phase I • X :/?-galactosidase = repressor • X • HOURS Figure 19.BIOLOGICAL ORDER repressor is is present. This the first phase. The by heterozygote of type (i~). according to the hypothesis. the synthesis of the enzyme gene If. is the second phase. however. an inducer is added. the female cytoplasm i~. being constitutive devoid of repressor. Something seems to be wrong. as soon as the gene z+ enters the female. Synthesis of /3-Galactosidase II by a Heterozygote. then found that the rate of enzyme synthesis decreases and reaches a zero value (Figure 19). This corresponds to constitutivity. thanks to the gene by the male. But the male has introduced also the inductive gene i+: is a repressor synthesized. the synthesis of galactosidase is followed long enough. it is [48] . continues. i+. and the synthesis of galactosidase stopped. and ^-galactosidase synthesized. In the zygote type II. shown in the preceding figure has been formed the transfer of the male genes i+ z+ into the i~ z~ female. A repressor has really been formed. however. is The is female.

And is it is quite natural that the synthesis of the contingent enzymes acids. genetic analysis has thus revealed a number of important The genes z. Thus. determines the constitutive as opposed to the inductive type of synthesis of both enzymes. lactose is a carbon source and a source of energy. The Ways of Repression Repressor. z and y. the bacterium stops synthesizing methionine synthase. One. It is called a regulating gejie.THE FUNCTIONAL ORDER If an inducer is tion of enzyme starts added after the synthesis has stopped. the control OF enzyme syntheses. Let us consider the enzymes responsible for the synthesis of the essential building blocks and especially of amino Amino acids. the hazards of to its The food daily life. ^-galactosides. The repressor has not yet been identified. is the biosynthesis of amino acid arginine. It is part of the food. They are called structural genes. But a number of data are in ^ favor of the hypothesis that it is a ribonucleic acid. It has already is methionine been stated that if the amino acid provided. Things happen as if the gene i+ controlling the inductive state corresponded to the i~ to the loss of the synthesis of a repressor. corepressor. y. For the bacterium. the small sector of the bacterial chromosome is that controls the synthesis responsible for the metabolism of lactose. This general. such as glycerol or pyruvic acid. and inducer. The bacterium may be fed with other organic compounds. of galactosidase and permease. the enzyme responsible for the last step in the biosyn- thesis of methionine. and a mechanism the known to be quite few cases have been studied extensively. Two carry the information for the synthesis. of the enzymes generally of more and of them. respectively. The product of enzymatic activity stops the is synthesis of the enzyme. aporepressor. i. that a bacterium depends on The bacterium obviously cannot afford find may manufacture a large number of useless enzymes. regulated. the mutation i+ ability to form a repressor. composed of three units. The facts. and i are independent units of function and therefore are different genes. hence to a constitutive situation. The other. the producanew. for In a example. [49] .

histidine and tryp- tophan. If supplied. The Biosynthesis of the Amino Acid Tryptophan. ® ® anthranilic acid ® (D tryptophane shikimic acid TRYPTOPHANE SYNTHETASE Figure 20. if an excess of arginine is produced.C. NH. Thus. such as nucleic bases. specific.BIOLOGICAL ORDER medium devoid arginine is of arginine. the synthesis of the enzymes responsible for the synthesis of tryptophan is not altered in the derepressed strain. In a synthetic synthesized by a medium devoid of tryptophan. repressed. and this excess tryptophan is excreted. Only the case of tryptophan will be discussed here. The produce much more tryptophan than needed. it is possible to isolate mu- tants that are called derepressed. Let blocked by a repressor. the bacterium synthesizes arginine. whereas it is inhibited the tryptophan enzymes bacteria [50] . If tryptophan is is is added. This repressor on the tryptophan enzymes and not on the synthesis of any other system. serine HO— C— C— COOH H I NHg COOH A. is and less arginine will be produced. the synthesis of is not blocked any more by tryptophan. In these mutants. HO- COOH /^ ® n indole C — C I COOH Y OH °" <d) NHg—••B. the synthesis of the enzymes involved in its synthesis is specifically repressed. the enzymes cease to be produced. If tryptophan is added to the medium. The same mechanism operative in a number of biosynthetic pathways. that is. the synthesis of the enzymes us admit that the synthesis is is stopped. as shown in Table III. The end product of the chain of biosynthesis blocks the synthesis of all the enzymes involved in the chain of arginine biosynthesis. it is known to act only From the original repressed bacterium.D. tryptophan system of enzymes (Figure 20).

c o D- u 4-> to (U c W c o c « Oh N c O I-H Cl- H c 'M o .

unable to manufacture the repressor. The without effect on the enzyme level in the derepressed strain.BIOLOGICAL ORDER Let us admit that the original wild type which is able to manufacture the repressor. is A obviously produced by the R+ gene. allowed to conjugate. Table IV. which dominant over the R~ gene (Table IV). Enzyme Synthesis as Affected by Tryptophan in Repressed and a Derepressed Bacterium and in the Heterozygote. the synthesis of the tryptophan repressor is enzyme is blocked. and that the derepressed bacterium carries a mutated gene R~ try. In this zygote is thus formed. The hetero- zygote behaves "repressed" is like the repressed strain: the character dominant. possesses the gene R+try. a The times derepressed bacteria contain three to seven the more enzymes than is repressed one. A and R~try genes. Without tryptophan . carrying both R+try heterozygote. R+ and R" bacteria are in the normal strain. whereas it blocks addition of tryptophan enzyme synthesis in the repressed one.

When growing in a medium containing glucose and lactose. but it expense of the reserves that the bac- A corepressor perhaps continues to be pro- has less affinity for the aporepressor than the product of glucose metabolism. The theory is that a product of the metabolism of glucose acts as a corepressor \'hich combines with an aporepressor. But is an excess of /^-galactosidase as a result synthesized. Lactose competes for the aporepressor with this hypothetical corepressor of low affinity and thus prevents is the formation of the repressor. But at the when it is used up. no /^-galactosidase start. consequently. The synthesis of the enzyme respon- for the metabolism of lactose will be repressed. In view of the relatively rapid adjustment of enzyme synthesis to the need.THE FUNCTIONAL ORDER Whether the tryptophan has been manufactured by the bacterium or pumped from the medium into the bacterium. is produced by the z+ bacteria. is glucose is used entirely. Things happen as if in the presence of glucose a repressor has been produced. A corepressor will be formed and. When it has disap- peared as a result of the cellular metabolism. an excess of glucose produced sible of its activity. a corepressor not produced or is is not available. duced. If The up if latter are used up in the metabolic processes. How does the aporepressor-repressor hypothesis apply to the regulation of enzymes involved in the metabolism of the carbon and energy source? When lactose is present as the sole carbon and energy source. the bacterium "needs" galactosidase in order to grow. it.8-galactosidase synthesized and functions: lactose is hydrolyzed into glucose and galactose. provided it possesses the corresponding genetic How does lactose induce the simpler to consider teria are first synthesis of the enzyme? It is perhaps bac- the inhibition of the synthesis. the synthesis of the enzyme can Why first then lactose necessary? Glucose has been used up in the phase of the bacterial development. but only after a negative phase that lasts around 30 minutes. The . metabolism continues terium has stocked. a repressor. [53] . Carbon and energy source. As long as this repressor is present. the result is the same. no ^-galactosidase first. Glucose is used up and the synthesis of /^-galactosidase starts. is is synthesized. and it starts synthesizing information.

no enzyme for its utilization is produced." This con- [54] .BIOLOGICAL ORDER this hypothesis implies that the repressor is is unstable and that its half-life short. the enzyme is useless. The growth rate of the constitutive bacteria is obviously lower than the growth rate of the adaptive ones. Anyhow. enzyme will not express itself if a repressor How does the repressor repress? Inductivity and con- stitutivity have to be considered anew. the bac- show that the constitutive bacteria contain about 30% more enzyme than the inductive ones. metabolite. In both cases. This the fact that part of the building blocks is is obviously due to utilized for the manulactose lactose facture of a useless enzyme. the When When enzyme is synthesized and the synthesis adjusted to the need. The structural gene carrying the information for the synthesis of a given present. insensitive to the whether or not lactose Estimates present. present. a preliminary attempt ization seems to be necessary. absent. When into a adaptive and constitutive bacteria are inoculated together of lactose. The effector of the regulatory mechanism is a repressor. is In the cases studied so far. is controlled by the "level" of this "Level" here means intracellular concentrawhether the essential metabolite is extrinsic or endogenous. the synthesis of the enzymes necessary for the synthesis of any given essential metabolites essential tion. Enzyme synthesis is of course limited by something. let us say a few minutes. in a medium devoid of lactose. Before any further discussion. Moreover. It at a general- might be stated that the synthesis of the enzymes required for the metabolism of the energy source is controlled by the over-all available energy sources. some others because is is the "operator" (explained in the next section) repressor. The sidase question will immediately be asked: How does /?-galacto- behave in a constitutive bacterium? Some bacteria are consti- tutive because they do not produce a repressor. is is Evolution has thus ended in a remarkable device. let us say by the availability of amino acids. inductivity "dominant. the constitutive bacteria are out- medium devoid grown by the adaptive ones and disappear. owing to natural selection. How does the repressor act? The is operator. terium contains yff-galactosidase.

desiraated by the letter The dominant the inductive + would have the o'' the constitution i+o+z + . and the constitutive gene i~. elaborate series of experiments has led to the conclusion that a when a bacterium mutates from recessive to dominant constitutivity. despite the presence of the inductive gene i + An hypothesis is that the gene produces repressor. The regulating gene produces — or does not produce —a specific repressor. no enzyme is formed in the absence of an inducer. the other i^z + In the absence . is dominant. Thanks to recombination experiments. specific gene is involved which has been called the operator and o. a The and the expression of the structural gene z+ of the chromosome i~z+ is blocked. 0+ gene being stitutive would be repressor. The func- tional unit formed by the operator gene and the structural gene has been called an "operon. . and next to it the regulating gene i. instead of being recessive.THE FUNCTIONAL ORDER elusion is two sets of based on the consideration of a heterozvgote containing chromosomes. The inductive gene i+ is said to be dominant. despite for constitutivity. The problem of the site of action of th. recessive (Figure 21). in position cis (Figure 22). The gene i+ imposes the inductive situation. It "sensitive" to the repressor. Thus. When constitutivity is dominant. that is. But affects only those genes located on the same chromosome. An extensive study has revealed that in some mutants constitutivity. the heterozygotes i + z + i~z+ manufacture the /3-galactosidase in the absence of inducer. of an inducer. The repressor produced thanks to the gene i+ acts on the "constitutive" chromosome carrying the constitutive gene i~ or on enzyme-forming system produced by the "constitutive" the chromosome.e repressor was solved by the discovery of a new category of mutants. despite the presence of the constitutive gene i~." The repressor formed by the combination of the aporepressor and [ 55 ] . no /?-galactosidase. The dominant cono"^ gene being "insensitive" to the was found it that the mutation o+ ^ affects not only (for the the gene z (for the /?-galactosidase) but also the gene v permease). i~o'^'z . no enzyme. The structural gene contains the informa- tion for the synthesis of the specific enzymes. the presence of the gene i~ responsible i+ is produced. one i + z+. Close to them is the operator gene o. it is known that the genes y and z are adjacent. .

dominant but only for the gene y+ located on the same The gene z+ is located on a chromosome carrying the wild-type operator o+ and is not active. is Inductivity dominant. The Operator Gene. No enzymes produced by the heterozygote in the absence of an inducer. located on the same both permease and iS-galactosidase are produced is in the absence of an inducer. 3. Constitutivity as o'' chromosome (position cis) . despite the presence of a repressor.BIOLOGICAL ORDER 1 - Inductivity i+ dominant operator + sensitive to repressor no/3 -golactosidase no permease 2ond3-Constitutivity operator 0^ insensitive I- dominant to repressor 2- 0'^/Z-*-y* in c/s fi -galactosidase present permease present 0VY+ no /3 inr/5 permease present 0<^/Z + in trans -galactosidase ^ •yj-galactosidase Opermease x repressor Figure 1. The operator are is of the wild type (o+) that is sensitive to the repressor. 21. i5e^ . are active: genes z+ and y+. The chromosome the constitutive operator o". Constitutivity as is dominant. 2.

for example.THE FUNCTIONAL ORDER of the corepressor would act on the operator gene. and enzyme synthesis is repressed. When the repressor is present. When no repressor is present. the operator gene is in position go. The structural genes z+ and v"^ "manufacture. the inducer. the operator gene o+ is in position go." respectively. Lactose. competes with the corepressor and prevents the formation of the repressor. When the inductive bacterium i+ is o + the operator is sensitive to the repressor. The Operon. When a repressor is present. and the question must be asked: How does the operator gene act? We have every reason to believe that the operator does not send messages to the cytoplasm. The repressor is produced from an aporepressor synthesized by the gene i+ and a corepressor that is a product of metabolism of the energy source — glucose. W' hen the repressor is absent. produce the enzyme-forming system. One hypothesis is that it acts by inducing a [57] . . the operator gene is in position stop. and the two structural genes z+ and v+ are "blocked" and do not OPERON regulating gene operator structural genes + Y+ ^____^ ~ ~ enzynrie forming systems /3-galactosidase permease co-repressor lactose metabolism Figure 22. /3-galactosidase and /3-galactoside permease only in the absence of the repressor. and the structural genes are active. There is a new problem now. the operator gene is in position stop.

however. All the members of a zymon work in harmony. is Co-ordinated control. if all the individual genes are linked and belong to the same operon. But the enzymes are there. and the activity of the already existing enzymes is depressed or stopped. the synthesis of the re- enzymes is repressed. but nothing more is known at the present time. A [58] . or first blocks. This corresponding to a right. for the genes of the is all tryptophan zymon. When the output ex- ceeds the consumption. And the activity of each enzyme depends on the activity of all the members of the zymon. This is zymon are some- the case. the genes zymon are not linked. It simple device. This would be the the genetic information for the zymon that is a unit. The study of bacteria has revealed the structural genes carrying the information corresponding to one times linked. Sometimes. acids. is When excess. the functioning of the enzyme of the biosynthetic path- way. Thus a and more sponsible mechanism controls the production of amino generally of essential metabolites. inhibition of enzymatic activity. The zymon is a functional whole. This conto exist in the arginine dition is known dual and the tryptophan zymons. One enzyme acts on the product of the activity of the preceding one. and if they continued to function. The most efficient and economical system of control would be a single system controlling the synthesis of case if all the members of the zymon. The other enzymes involved synthesis are activity of the in the particular chain of bio- consequently deprived of their substrate. Let us consider first the synthesis of an essential metabolite: tryptophan. happens that this accumulation is prevented by a very The end product of enzymatic activity inhibits. Preclusive and corrective feedback mechanisms. that the need. Refinement of regulation. is. the accumulation of the end product would continue. The group of enzymes that for the synthesis of a given essential metabolite or responsible more generally for a given sequence of reactions will be called a zymon.BIOLOGICAL ORDER functional change in the structural genes. the end product of a biosynthetic process a produced in repressor is formed which blocks the synthesis of the corresponding enzymes. But will be found. and the zymon is reduced or blocked (Figure 23). it And for this no explana- tion has yet been found. for example.

THE FUNCTIONAL ORDER OPERON structural genes reguloting gene operator L^^' enzyme forming systems Qd O/i Oy 6^ enzymes .

zymon. the repressor is one of enzymes different from the system submitted to the repression. The corrective feedback deals with the synthesis of an essential the end product of the metabolite. counteraction. 318-356. the bacterium. 26. 1961. no single molecule or group of molecules can be held responsible for *The reader interested in the control of protein synthesis and in the problem of regulation is referred to the excellent review of Jacob and Monod. or one group of persons. microorganism. one person. a repressor is the synthesis of ^-galactosidase. and to Jacob and Wollman. has been compared to a factory. [60] . The of all organism. In order to survive. it must adjust its enzymatic equipment according to the nature of its food and to the nature of its needs. And also. many more systems have to be analyzed before any attempt at a generalization can be made.* the tryptophan the end products of a system of Conclusion The study of enzyme synthesis has thus revealed that the cell is endowed with an elaborate dual mechanism which controls the activity and the synthesis of enzymes. The repressor prevents present. In the first case. Some in a readers may be tempted to ask the question: in a Who commands bacterium? Obviously. Things are different when the energy source glucose that is is is concerned. When produced. The organism has to cope with the variation of the environment. In a factory. Sexuality and the GeJietics of Bacteria. Academic Press. the corepressor is activity of the enzymatic system submitted to repression. In the other case. MoL BioL (1961). thanks to a corepressor a by-product of glucose metabolism.BIOLOGICAL ORDER negative feedback mechanism: in this case the control is corrective. the machine must be regulated in such a way that each one of the four nucleic bases and each one of the twenty amino acids is manufactured in just the right amount and proportion. The activity of the enzyme re- sponsible for the metabolism of one energy source prevents the synthesis of a system involved in the metabolism of another energy source. /. directs the activity the others. New York. the preclusive feedback with the metabolism of an energy source. Obviously. namely. or preclusive. but it is in fact a preventive. This mechanism could be considered as a feedback. to the last issue of the Cold Spring Harbor Syinposia on Quantitative Biology (1961). 3.

685-691. Angew.. eds. G.THE FUNCTIONAL ORDER the harmonious dynamic cellular balance. The enzymes and the comparator. A Syinposium on the Chertiical Basis of Develop7Hent. Repres(1958). Symposium on the Regidation of Cell Metabolism. Induktion. itself is metabolism the result of the interaction of and of the dual feedback mechanism. W. The Johns Hopkins Press. Wolstenholme. This formula should take on its full significance. it is simply suppressed. an essential part is function to compare the result of the \vork done by the machine with the information fed to the computer. ed. The functional order is the result of the interplay of the hereditary material. Structures and functions are the is complementary aspects of of order. sion.. New McElroy. There is no such thing as a comparator in a microorganism. E?izy??!es: U?iits of Biological Structure and Function (Henry Ford Hospital. Evolution has necessarily selected the fittest types. Biosynthese eines Enzyms. of the enzymes. Che?me. The living organism has been defined as an independent unit of integrated structures and functions. The organism a functional system REFERENCES Gaebler. When the work does not fit. of the metabolism. E. International Symposium). J. Boston. W. pany. Its food. Baltimore. and Glass. (1958). Instead. A now specific function can only be the expression of a specific structure. the struggle for life. Academic Press. H. Brown and Com- [61] . 11. eds. C. and O'Connor. Information. as proyided by the environment. O. D. (1959). M... B. each organism takes part in a compe- tition. life. The comparison a functional one. Ciba Foundation Little. Alonod. in which the functional values of the is different types are compared. York. (1956). is In a computer..

one discovers en- able to reproduce their kind is which are about 250 A in di- ameter. which to organize these enzymes a The ultimate unit of integrated structures and size. Two systems of order are then intertangled. Their volume therefore 8. functions must necessarily possess a minimal As matter of fact. They are strictly dies. the diameter of the smallest microorganism capable of independent reproduction When tities is about 5. The cell must at its disposal some 2. to grow. a living cell. as such. the is unable to metabolize. It must therefore have possess the genetic information for the synthesis of these in 2.000 enzyme species in order to secure energy and to synthesize all its building blocks. the cell/virus system survives and multiplies.VIRAL FUNCTIONS: ORDER AND DISORDER Introduction An kind. Viruses can develop only inside intracellular parasites. Viruses are able to reproduce their kind. These entities are viral particles. A cell infected by a virus very often Sometimes. The viral order is [62] .000 in the enzymes and some space proper organelles. tion organism is an orderly system of integrated its structures and functions able to metabolize and to reproduce The microorganism or the cell is the ultimate unit of integra- and reproduction. however.000 times smaller than that of the smallest microorganism. viral particle.000 A. Yet. Ulthuate here means smallest. one studies the living \vorld. Viral multiplication thus poses a certain number of problems. however. They contain proteins and nucleic acid and exhibit a constant and welldefined structure. and to undergo binary fusion.

viral functions. which means a unit of virus. the poliovirus. Various aspects of viral order. will be taken as a proa virion. The virion is a polyhedron. modern. Protein coat Subunit Nucleic acid Figure 24. and to be terminology. It has been crystallized and its crystals studied by X-ray diffraction. The most general idea of a viral particle can be derived from the study of one of the simplest and smallest viruses. It is composed of a protein shell en- closing the nucleic acid (Figure 24). Schematic Representation of a Viral Infectious Particle (Virion). The virus of poliomyelitis. and cell/virus interactions will be discussed in this chapter. In order to simplify things. or cancer when the mammalian cell is involved. As a consequence. the infectious viral particle will be called virion. made of subunits. the cellular order is perturbed. \^iRAL order: the infectious viral particle. X-ray diffraction shows that [63] . according to recent to spare words. or virion. arranged in an orderly fashion. totype of This virus has been obtained in a high degree of homogeneity and its chemical constitution established. the capsomeres. The The genetic material (nucleic acid) capsid is is folded and enclosed in a coat or capsid.VIRAL FUNCTIONS: ORDER AND DISORDER superimposed on the cellular order. and the results may be as different as "lysogeny" when bacteria are concerned.

the virion is metabolically inert. acids. its The is. viral genetic material. These conditions exist only inside a cell.600 amino acids. Even if some of the proteins are endowed with a potential enzymatic activity. the maximum number of amino acids that can be organized in the viral nucleic acid is 1. is number of protein It species.700. which is the critical living mass. No is other "biological" particle or a large organelle yet known which composed of number of asymmetrical subunits exhibiting relations of symmetry. This type of structure seems to be able to form only a viral coat and nothing else. built of subunits. that itself. to synthesize the specific viral proteins and must necessarily be in an environment where building blocks and energy are provided. The in structure of the viral it infectious particle is extraordinary the sense that is is unique.600 amino of smaller molecules. The molecular weight of of the virion is approximately 6. It is built up out of 60 structurally equivalent asymmetric units of approximately 60 A diameter. the conclusion was reached that a sequence of three nucleotides was probably necessary to take care of one amino acid. in order to express potentialities. The capsid can only be and is in fact built of subunits. This type of structure is clearly seen in electromicroscopic photographs of many viral species. The genetic material of the virion therefore cannot directly build a molecule composed of 49. The nucleic acid is composed of about 5. which the protein represents about 62%. is but it can build a number Each capsomere in fact made up of approxi- mately 620 amino acids. Therefore. The protein coat of viruses or capsid close contact in a form a is actually made up of subunits or capsomeres. to duplicate The essential function of the protein viral coat is to provide a cell to cell means for the transmission of the genetic material from [64] . they roughly spherical shell of about 300 A diameter.200 nucleotides. An entity containing a small five. cannot secure energy and cannot synthesize its building blocks. If 60 protein subunits of that size are packed together in way consistent with icosahedral symmetry.BIOLOGICAL ORDER each virion possesses an icosahedral symmetry. In the discussion of the code. the nucleic acid 28%.7 X lO^*. let us say one to unable to do much work. The protein of the capsid contains about 49.

let us say poliovirus. it is The proof of the effectiveness of the infection the production virus is by the infected cell of a new generation of virions. The genetic material is RNA in some viruses.000 particles. and which is an essential and clear also that the genetic material of information (a) for the synthesis of the pro- teins of the viral coat. This is the so-called vegetative phase of the viral are not life cycle. A virion never contains both nucleic acids. the synthesis of cellular structures a well-ordered process. properly handled and not altered. viral proteins are synthesized. subject to a regulating system? If a viral regulating system does exist. Moreover. reproduction from the genetic material only specific feature of viruses. They -appear around the third hour. a An elaborate system of repression and induction based on feedback mechanism controls the balanced as synthesis of enzymes. and (b) for autonomous reproduction. pathological particles of Are viruses. and DNA. in others. Every every micro- RNA nor a organism without exception. Remarks on the control of viral functions. Approximately 6 to 8 hours after the infection each cell contains about viral 100. how do viral and cellular functions interact? The study of bacteriophage should allow us to answer these questions. It a virus has to possess the is is something which neither a cell microorganism can accomplish. The nucleic acid of a If number of viruses has been if extracted and purified. infective particles. [65] . the cell damaged and dis- integrates. In a normal. DNA cell. and the genetic material is multiplied. infect cells. among which only about 500 is are infectious. reproduced from its genetic material only. finally As a result of viral multiplication.VIRAL FUNCTIONS: ORDER and a protection against the adverse conditions AND DISORDER in the encountered outer world. An organized infectious entity possessing only one type of nucleic acid can only be a virus. The may The virus. like normal cellular constituents. and thereafter will increase in number. produced during the first phase. When a cell is infected by a virus. At the is cellular level. noninfected is cell. a viral infection the penetration into a cell of the genetic material of a virus. Mrions. possesses both types of nucleic acid. Viruses are sometimes considered cellular origin.

the bacteriophage becomes external attached to the bacterium (Figure 26). But when the tip of its tail into contact with the bacterial wall. (a) a contractile outer sheath. composed of is a ball of DNA surrounded by is a protein coat. Figure 25. tail The is contracts. is The synthesis of phage protein initiated. (b) an internal tube. The bacteriophage is more complicated than the poliovirus. and (c) tip. The genetic material through the tube into the bacterial cytoplasm. It has a so-called "head" and a so-called "tail" (Figure 25). The head is a hexagonal particle prism. The internal tube penetrates through injected membrane a inside the bacterium. and 5 minutes after the [66] . The tail a long cylindrical structure. Schematic Representation of a Bacteriophage Particle. sheath of the the An enzyme present at the extremity of the phage tip attacks the bacterial wall. The comes bacteriophage is not motile.BIOLOGICAL ORDER Bacteriophage Life-cycle of a virulent bacteriophage. which composed of a. All within minute. this happens Then the so-called vegetative phase takes place.

After 12 to 20 minutes. The bacterium is lysed and liberates some hundred [67] . The chromosome c is disintegrating. the is tail appears \yhere it should appear. bacterial chromosome has disappeared. 26. This endolysine depolymerizes the mucopolysaccharides of the bacterial wall.VIRAL FUNCTIONS: ORDER infection the genetic material AND DISORDER of the bacteriophage starts multi- plying. and the autonomous multiplication of the phage genetic material is initiated. has injected genetic material into the bacterium. The bacterial 3. a peculiar phage protein has been produced. (b) 2. condensed into and the subunits of the head are assembled around the ball is DNA <v/w» — genetic material of the bacteriophage endolysine • phage proteins Figure 1. The \vill infectious particle formed. Finally. Its building blocks (nucleic bases) will be incorporated in the phage genetic material. and phage particles are formed. A lytic enzyme (endolysine) bacterial wall will be will be vegetative bacteriophage. thus liberating the bacteriophage particles. The it Life Cycle of a Bacteriophage. by a process that is not understood. 4. of the In the meantime. The produced by the hydrolyzed and the bac- terium will l)-se. The proteins are organized around the folded genetic material. Beginning of the vegetative phase: Phage proteins are produced. the be localized at the tip endolysine. the bacteriophage particle has attached onto the receptive its Infection: (a) bacterium. of nucleic acid. More phage material has been synthesized. the phage a ball. \yhich in the particle tail.

BIOLOGICAL ORDER bacteriophage particles. called viriile?it phages. called temperate phages. Others. Some bacteriophages. are multiplied only by the lytic cycle. The so-called "lytic cycle" of the bacterio- phage has been completed. may evolve in two directions. / c .

The lysogenic bacterium multiplies a specific structure: the prophage perpetuate to (etymologically. immunological specificity. shape. 1953). etc. the bacterium grows and divides. it dies. CDnon-lysogenic b ( [ >*/ Productive infectio i :'c^ phase ^g^ Vegetative [^~:cc:<-J CHD >{Em^ Figure 28. which might differ by a number size. A number of experimental data have led to three essential con- clusions (Figure 28). When this potentiality is not expressed. it will perpetuate the potentiality to produce B. [69] . of properties such as host range. Diagrammatic Representation of Lysogeny (after Lwoff. When infected with B. us say A or B. "before the phage"). A given bacterial species can be infected by let a of different strains of temperate bacteriophages. When it is expressed. The number prophage.VIRAL FUNCTIONS: ORDER possesses AND DISORDER and perpetuates the potentiality of producing bacteriophage particles in the absence of infection. A given bacterium infected and lysogenized by the potentiality a temperate bacteriophage A will produce bacteriophage A. The bacteria that perpetuate the possibility of producing bacteriophage in the absence of infection are called lysoge7iic.

itself after an infection. How is it possible that one is dual behavior? This But there prophage. and the vegetative phase culminates with the organization of infectious particles. It carries the information for the production of bacteriophage particles. according to the bacterial species. is always attached to the same site. is homologous chromosomes takes place. [70] . the genetic material of the bacteriophage bacterial rial attached to the chromosome. 2. 3. originated from the Prophage and vegetative phage. the its chromosome 2 multiplies at own pace. of the bacterial chromosome. has chromosome. bacterial in favor of the hypothesis according to which the genetic material of the phage has. Cytologists know that during a certain phase of the maturation of the sexual cells. behaves differently from the vegetative genetic material. the bacterium is lysogenic. The prophage that is. The nature of unknown. every 20 to 50 minutes. viral During the vegetative phase. admitted that the attachment of the of the bacterial viral genetic material on a specific site the consequence of a correspondence of structure of the chromosome two is interacting nucleic acids: the bacterial one and the viral one. or locus. duplicates itself alongside the bacterial chromosome. When. In a lysogenic bacterium. The phage does not express its potentialities: no viral proteins are to be found in a lysogenic bacterium and of course no infectious particles.BIOLOGICAL ORDER 1. etc. the prophage. During chromosome is autonomous. That a given prophage site is is able to reach and recog- nize a specific chromosomal something strange. The prophage There is is the genetic material of the bacteriophage. it is. The genetic mate- of the bacteriophage. long ago. a pairing of stick The two chromosomes regions. This correspondence. the phage medium. as prophage. The receptor. temperature. another difference between vegetative phage and vegetative viral The chromosome expresses its potenpro- viral proteins are synthesized. It duplicates itself every or 3 minutes. nature of the the vegetative phase. A given prophage only one prophage per bacterial chromosome. tialities: is and the same structure exhibits such a one of our problems. submitted to bacterial control. whether it is homologous or complementary. to each other at their homologous is the forces that intervene in the process it is In any event.

however. the lysogenic bacterium does produce phage. the answer. But thanks to the presence of the prophage the lysogenic bacterium exhibits another property. the genetic material of phage A does not multiply and of bacteriophage. and each one has liberated some [71] . These strains are called inducible. In other can be induced at will in practically all of the population. Other types us say B.to 10~" Induction. is diluted out in the course of bacterial is multiplication (Figure 29).VIRAL FUNCTIONS: ORDER AND DISORDER defined as a bacterium that perpetuates produce bacteriophage in the absence of infection. spontaneous because \e do not know cause. Then within 10 minutes. not genetically related to bacterioif prophage A were not cytoplasm of the lysogenic bacterium carr\ing prophage A which prevents specifically the vegetative multiplication of the genetic material of phage A. the production of bacteriophage cannot be modified. The The simplest hypothesis study of lysogeny should provide is that viral functions are not expressed in a lysogenic bacterium because they are blocked by a repressor. ImiMumty. Let us assume that our bacterium has been lysogenized by a temperate bacteriophage A and therefore perpetuates a prophage A. The per bacterium per generation. Viral functions are not expressed. Phage proteins are not produced. the so-called im- munity. The superinfection by the homologous bacteriophage A does not result in a vegetative phase. Moreover. From time to time. frequency of spontaneous production In some lysogenic strains. the is the so-called spontaneous its phage production. phage A present. This strains. something can multiply vegetatively. All the bacteria have been lysed. Thus. but behaves as an inert particle. The genetic material of the infecting it bacteriophage A is injected into the lysogenic bacterium. Inducible lysogenic bacteria are irradiated with a suitable dose of ultraviolet light. the bacterial population disappears. a lysogenic bacterium perpetuates the potentiality to produce phage. The bacteria continue to grow for about 45 minutes. This let immunity specific. probability that a lysogenic population will produce infectious particles varies with the strains from 10 ~. and we have to know why viral functions are sometimes the A lysogenic bacterium was power to expressed and sometimes not. just as must be present in the Here again a specific repressor of viral functions can be visualized.

and One of the daughter bacteria (3) contains only the prophage. by an irradiation with ultraviolet The vegetative phase of both prophages and the super- infecting phage are initiated. This lives in is harmony with its prophage. most extraordinary. The problem of the control of viral development is thus posed in the most dramatic way. of i V y- (p) or corresponding autonomous D. A bacterium The prophage behaves as if it suddenly. the lysogenic bacterium has dies. gous phage. 4-5. for example. the vegetative phase of the bacteriophage initiated.N. is induced.BIOLOGICAL ORDER 100 infectious particles. A. as a result of an irradiis were a normal bacterial gene. The genetic material of the superinfecting phage does not initiate is the vegetative phase. 29. lysogenic bacterium produces bacteriophage and to the irradiation. does not divide. previous been infected with a mutant of the homologous phage toward which the bacterium is immune. The superinfected bacterium light. and the i^r prophage D. the mutant will develop together with the prophage. is The lysogenic bacterium superinfected with a mutant of the homolodiluted out at each division. homologous superinfecting phage Figure 1-3. [72] . And ation. A. Immunity.N. If. and infectious particles of both types will be produced.

This These data are consistent with the hypothesis that in the lysogenic male bacterium the expression of the viral functions is blocked by a cytoplasmic repressor. an inducible prophage penetrates zygotic indnctiov. All these agents are known to be mutagenic and oncogenic. the vegetative phase is initiated. the vegetative phase initiated. epoxides. Ultraviolet not the only inducer. and viral functions are How is could the repressor act? The antibiotic chlor- amphenicol known to block the synthesis of proteins. that is. The prophage A of Escherichia coli K12 is located on the bacterial chromosome close to one of the genes controlling the utilization of galactose. induction of a lysogenic bacterium is not homologous to a mutation. also inducers light has AND DISORDER is an inducing effect but A number of physical or chemical agents are — among them. cancer-inducing. If only the female if a lysogenic. a fact that is both male and female are lysogenic. female is lysed. Anyhow. phage development is induced when. If bacterial [73] . and ethylenimines.VIRAL FUNCTIONS: ORDER Inducing agents. nitrogen mustard. the level of the repressor decreases is below a given threshold. nothing happens either. As a consequence. Thus. the theory that the inducing agents depress or block the synthesis of the repressor. and the is genic male has penetrated into the nonlysogenic female. as a result of conjugation or zygosis. repression ceases because the cyto- plasm of the female expressed. is devoid of repressor. If conjugation. things are quite different (Figure 30). or 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hich has a greater affinity for the repressor. The fact that inducing agents are mutagenic is not a great help: organic peroxides. But the fact that inducing agents are able to initiate malignant growth Perhaps induction will help us to understand is rather exciting. cancer. X rays. which enters into the female at about the 25th minute after conjugation. gamma rays. or a more stable repressor. a A tures noninducible prophage could be prophage which manufaclater. but the reverse is is certainlv not true. This problem will be discussed Zygotic induction. But a lysogenic male mates with nonlysogenic female. Since the prophage is no longer repressed. as the As soon prophage attached to the chromosome of the lysobacteriophage particles are produced. more repressor. nothing happens after is worth while noticing. When the prophage of the lysogenic male enters the nonlysogenic female. into a nonlysogenic cytoplasm.

Prophage during Bacterial Recombination. If this interpretation correct. zygotic induction. The vegetative phase is not initiated. — phage proteins X repressor Figure 30. Upper row: The lysogenic male (1) (repressor present) injects its chromosome and prophage into the lysogenic female (2) (repressor present). It could be that the viral function necessary for the onset of the is vegetative phase is the synthesis of a protein. it should be applied to infection and lysogenization. zygotic induction does not take place in a large fraction of the zygotes.BIOLOGICAL ORDER phage D.N. [74] . The vegetative initiated: this conjugation takes place in the presence of chloramphenicol.A. Lower row: The some and prophage phase is lysogenic male (4) in a is (repressor present) injects its chromo- nonlysogenic female (no repressor).

and the bacterium viral now lysogenic. It continues to we shall try to understand why produce the repressor. Something irreversible has happened.) acts in favor of lysogenization (Figure 31). If a protein is is produced is the vegetative phase initiated. the viral DNA duplicated. Thus. But.VIRAL FUNCTIONS: ORDER AND DISORDER fected Infection and lysogenization. does. namely. like any other genetic material. It is now a prophage. the viral site. The outcome of the infecon the genetic constitution of the temperate bacteriophage. material of a bacteriophage. DNA The it is to this receptor later on. If a repressor is produced first. is a repressor. A nonlvsogenic bacterium inby a temperate phage may either produce infectious particles and be Ivsed.) a low temperature (20°C. the expression of the viral potentialities is blocked. A high whereas temperature (40°C. A mutant ind^ was discovered which had lost its original inducibility [75] . and finally infectious particles are produced. and the process irreversible from More proteins are synthesized. and the fate of the bacterium. the behavior of tion depends the bacterium/phage system. favors the establishment of the lytic cycle. or become Ivsogenic. thanks to the correspond- DNA ence of structure with attaches itself a given chromosomal locus. After the seventh minute the decision made and cannot be modified. But with a given temperate bacteriophage. The genetic mutants. and it will be reinforced by the study of Mutations affecting inducibility. and the autonomous cytoplasmic viral is unable to duplicate itself. the experimental data point toward the conclusion that the response of the infected bacterium depends on a primary determining event. What It is this event? that antibiotics was found first which block protein synthesis. The low or high temperature acts only during the first 7 is minutes after infection. are controlled by extrinsic factors. may undergo muta- The wild-type A ind+ is inducible with ultraviolet light. The fraction of the infected bacteria that will be lysogenized varies effectively from less than \^c to more than 99%. such as is chloramphenicol. something then on. else. favor the lysogenic response. that the The a hypothesis event can be the synthesis either of protein or of first. tions. The repressor hypothesis needs of course to be reinforced. Proteins cannot be synthesized. prophage multiplies.

and infectious particles are produced. ^ phage proteins X repressor Figure 31. is.) in the presence of chloramphenicol.V. or in a bacterium irradiated with ultraviolet rays. The vegetative phase is initiated. The Fate of the Infected Bacterium.). ble with ultraviolet light Double lysogenic strains The mutant not inducihowever. and the genetic material of the phage reaches its chromosomal locus and becomes a prophage. a As with ultraviolet light. can be obtained that carry two prophages A: [76] . sensitive to zygotic indu<iition. although all other characters of the mutant were the same as those of the original strain. is The nonlysogenic bacterium 1-3: at infected: low temperature (20°C. X X •\AAAphQge A.BIOLOGICAL ORDER nfection 20°. 4-5: at high temperature (40°C. The first activity of the genetic material of the phage is the synthesis of a protein. or in first bacteria deprived of food. U. N. the vegetative phase is not initiated. starvation 40°. Chloromycetin. the repressor. X ^. activity of the genetic material of the consequence the synthesis of phage protein is repressed. lysogenizotion ' vegetative phase X X X ^ ^ X X X X V X ^ X X X X X X X y X X X D.

When a bacterium carrying prophage A ind+ is irradiated with ultraviolet light. Table V. Inducible .VIRAL FUNCTIONS: ORDER the inducible one. The noninductive mutation A ind" is dominant. the phage A ind+ cannot be induced. AND DISORDER these strains are When none of the bacteriophages develop. Variation of Inducibilitv. irradiated with ultraviolet light. it is induced and will produce phage (Table V). In the presence of A ind". and the noninducible one.

Ability to lysogenize . Mutations Affecting the Life Cycle.BIOLOGICAL ORDER Table VI.

and the is genetic material of the phage. Defective Prophages. . more of these proteins is impossible. And apparently.VIRAL FUNCTIONS: ORDER AND DISORDER Some are able to the synthesis of one or manufacture the proteins of the phage. is not multiplied. It seems as though the vegetative phase could be characterized by a sequence of events. In others. so far as known. the chromosome of the defective bacteriophage Table VII. in the absence of protein synthesis. Each event has to be completed so that the following one can be initiated.

Moreover. This happens also if the repressor balance is upset by inducing agents such as ultraviolet light. 1. The most important aspect of the reproduction of the viral genetic material has not yet been discussed. and they produce the proteins necessary for the unable to produce onset of the vegetative phase of their It own life cycle. has been admitted that the prophage does not enter the vegeta- tive phase If this because it is unable to produce some specific protein. Their properties seem to be controlled by their position in the bacterium. 2. the homologous prophage enters the vegetative phase. Let me repeat that the virulent mutants of a temperate phage are a repressor and are insensitive to the repressor produced by the original prophage. The synthesis of this protein may be blocked by a specific repressor. If a lysogenic bacterium is infected with a virulent phage genetically related to the prophage.BIOLOGICAL ORDER the autonomous if replication of the viral genetic material can take is place only the genetic material of the phage no longer exposed to the repressor. the behavior of two structurally identical viral chromosomes might be different. the virulent mutant. whereas the pro- phage multiplies. the chromosome of the superinfecting bacteriophage does not multiply. All the experi- mental data are thus in agreement with the hypothesis that autono- mous multiplication of a phage requires the manufacture of a protein. This happens when the prophage of the lysogenic male enters a nonlysogenic female where no repressor is present. which does multiply and which does produce these proteins. should induce the vegetative development of the prophage. In a lysogenic bacterium. the prophage is duplicated in harmony with the [80] . They develop despite the presence of a repressor. whether it is attached to the chromosome or to an episome such as the sexual F factor (Figure 32). Thus. hypothesis corresponds to the facts. Position is the fourth dimension of the prophage. Interactions of Cellular and Viral Functions Cellular control of viral reproduction.

The prophage is obviously subject to the system that controls the duplication of the bacterial chromoquestion some. is: What is this system? As a matter of exist? fact. that is. occurs every 20 to 50 minutes. the first Does such a system .VIRAL FUNCTIONS: ORDER cellular organelle to AND DISORDER one duplication which it is attached.

In one case. The enzymes for tryptophan synthesis may be produced while those for methionine synthesis are repressed. It happens that this gene galactoA. Thus the genetic material of a virus. A part of the receptor of prophage A the gene that carries the information for galactokinase (the kinase can be exchanged with a part of phage kinase is enzyme that phosphorylates galactose). the prophage behaves as In the other case. The prophage behaves is were a bacterial element. all The existence of such functions chromosome can only be postu- strong argument in favor of this hypothesis is that in a cell chromosomes divide not only at the right time but also at the same time. submitted to the cellular system of control. Obviously. The study of the behavior of the defective prophage after induc- tion has revealed the existence of a set of functions controlling the duplication of the viral nucleic acid. The reverse is also true: is cellular gene may be under viral command. of the induction and repression of enzyme synthesis. In a bacterium. some unknown factor commands the the duplication of the chromosomes. has revealed a different situation: the various individual systems controlling enzyme synthesis are inde- pendent in the sense that one may be blocked while the others are working. when a attached to a cellular organelle.BIOLOGICAL ORDER and persist in a prophage which is reproduced as nucleic acid and which can be perpetuated without expressing its potentialities. the A chromosome and as if it the prophage act as a whole. were a viral gene. nonlysogenic If Sahnojiella produces given somatic antigen lysogenized [82] . the gene galactokinase multiplies too. Are viruses able to take command of cellular functions? The reader realizes of course that the question would not be posed of a if a positive answer could not be provided. Viral control cellular functions. given chromosome. a bacterial gene behaves as This probably means that viral. A 10. and when this The gene galactophage A multiplies vegetatively. whether cellular or if it if it and the "foreign" nucleic acid it may carry constitute a unit and behave as a whole. The prophage the bacterial attached to a specific receptor site of is chromosome. a were a bacterial gene. so far as replication is concerned. now part of phage A. for the duplication of the cellular lated. The study of the functional order.

Thus a virus may be subject to the control system of a bacterium. and not in position trans (when they are in different structures). both for replication and for expression. turn produces another antigen The mechanism by which is proa phage 15 controls the synthesis of a bacterial antigen still mvsterv. Thus may be multiplied as genetic [83] . A much more fascinating situation has been disclosed A short digression necessary here. Things happen here as if the bacterial gene were obeying some viral mechanism of control. The gene is with galactogalactokinase and the prophage A are closely linked. dies. either as a result of an infection or as a consequence of an induction. then the enzyme galactokinase is synthesized in large amounts in the absence of an inducer. Strictly speaking.VIRAL FUNCTIONS: ORDER with given phage AND DISORDER a e 15. If the partners are properly selected. the defective prophage is derepressed. a large variety of heterozygotes may be obtained. This synthesis takes place only when the gene and the phage are in position cis (when they are part of the same structure). When a defective lysogenic bacterium is induced. the disease is not viral. but the bacterium nevertheless result of an infection. and the hypothesis that the genetic material of viruses has is evolved from cellular genes currently admitted. bacteria contain only a small synthesis of the amount of A is marked present. Both may be attached to the sexual factor F. A few hereditary. The defective prophage possesses some of the normal viral genes but behaves as a potential lethal gene (Table VIII). enzyme takes place only when an inducer But when phage A develops vegetatively. nonviral diseases of bacteria are known which are controlled not by a defective prophage but by genetic structures that have to be visualized potentially lethal genes. and a bacterial gene to the control system of a virus. Viral and noxviral diseases. as bacterial genes. and in 15. it stops producing the antigen 10. and thus transferred from the male to the female. The disease is not the and infectious particles have not been formed. It happens that in the absence of an inducer the galactokinase. perhaps a viral operator. An abnormal vegetative phase is initiated. Bacteriophage particles are not produced. diseases. kinase. a virus Viruses and cancer. Between these hereditary series of diseases and viral an almost complete intermediary steps has been disclosed.

BIOLOGICAL ORDER Table VIII. Inductive gene . Genes and Viruses.

But the question possible It is is asked: is What can originate from order? the only answer disorder. viral functions are repressed. the nature of the cellular modidue to the virus or provirus and responsible for malignancy is still unknown. must not cell the cell it infects. When may endowed with its the potentiality for producing virus infection. Yet it is interesting to consider the animal cell/provirus system. The cell original cell was normal. according to the nature of the one way could be cell and of the virus. in a order to be oncogenic.VIRAL FUNCTIONS: ORDER for AND DISORDER it cancer. cell it is is sometimes killed. It or the other. The cancer problem fication not solved so far. the cell modified by the virus. the bacterium is an is which is organism. clear that the superposition of two types of structural order may be responsible for functional disorder. something more: the animal cell/virus system is it multiplies and kills its now host. as a result of an alteration of the cell's environment. animal cell/oncogenic virus systems. these The transformed yet-unknown factors. The it infected survives. Biological order origin in primitive disorder. Whether cell has lost its sensitivity to or not the genetic material of the virus multiplies autonomously as provirus is not yet known. Darkness comes from had its light. not the oncogenic virus fact that specific repressors are a that kills the cancerous animal but the cell/virus system. Injected into an animal. As already stated. At least one thing is certain. and kill killed. cold from heat. When. But there is malignant. it is immune also be modified in morphology and physiology. low virulence for the It is. According to the Platonic concept. to control The known some viral functions opens the problem of oncogenic viruses. subject to the factors that control reproduction. It in the absence of infection. Structures and functions [85] . new way of approach to all things originate in their opposites. pleasure from if pain. The animal as if it were independent. may mean a high degree of pathoitself genicity for the animal. however. sometimes to super- survives. A And virus. In some of the Viral antigens and infectious particles are not produced. viral antigens and viral particles are produced. viral functions are derepressed. the normal cell a malignant the host is cell behaves in the dependent part of an organism.

Genetic control of viral functions. Burnet. VVollman. 1709-1714. Lysogeny. S. — Biochemical. Interscience Publishers. New York. The Viruses J. 92-111. eds. 1-39. E. E.. transduction and cancer genesis. viral diseases Lwoff. F. M. and Stanley. Control and interrelations of metabolic and of bacteria. (1958-1959). La sexualite des bacteries. F. New York. Masson. series 54. and Jacob.BIOLOGICAL ORDER are complementary in processes. New York. L. The Harvey Lectures. T. Monographies de rinstitut Pasteur.. F. University of Texas Press. Nature. (1954-1955). John Wiley & Sons. [86] . New York. Volume I. Press. Luria. Bacteriophages. pp. Paris. A. Biological disorder logical order. series 50. (1959). H. (1959). F. Biological. New York. and Jacob. Austin. WoUman. M. Academic and Klug. Academic Press. Academic Press.. pp. and Biophysical Properties. normal processes as well as in pathological is the complementary aspect of bio- REFERENCES Adams. AL. Jacob. Structure of poHomyehtis virus. Finch. 43-59. A. The Harvey Lectures. 183. General Virology.. W. (1959). L. (1959). (1959). 43-59. In Genetics and Cancer. (1953). E.

BIOLOGICAL ORDER AND ENTROPY Sources of Energy a An organism is a maciiine. When is the organism possesses chlorophyll. The organism performs oxidoreductions. or mineral sulfide. Max Delbriick has described an organism as a system of flux equilibrium which takes matter and energy from its environment. It is not coal. the reaction AH2 This is + /z Oo —> A + HoO respiration. The organism. may be written: AHo hydrogen donor (reduced If. The general equation as sugars or amino acids. the source of energy energy in this is light. case. for example coal. When the organism is devoid of chlorophyll. but the combustion The living machine. however. in order to operate. that causes the steam engine to operate. of coal. Aietabolism and syntheses consist essentially in the mobilization of the energy of the chemical bonds of the food. burns more-or-less complex substances: organic com- pounds such as hydrogen compounds such carbon monoxide. machine has to be fed something. It needs energy. the provided by the chemical bonds of the food. the hydrogen acceptor oxygen. -f B hydrogen acceptor (oxidized state) — is A oxidized + BHo reduced acceptor is state) donor for example. or even hydrogen. In order to operate. must also be fed. [87] .

and the free energy of reactions ( measure of the maximum amount of work — obtainable. In the reaction: .BIOLOGICAL ORDER Equations In any chemical reaction. one has to distinguish between the total heat of reactions ( — aH) a The free energy F gives aF).

" that is. the probability increases. namely. Boltzmann constant and D is the measure of the The state of rest which an isolated system tends to reach corresponds to a maximum entropy. and an increase of order to a decrease of entropy.38 X 10" ^'^ and P the number corre- "elementary complexions. 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is a measure of atomic disorder. to the jumps of the atomic system from one metastable structure to another. An increase of disorder corresponds to an increase of entropy. that 1. from a thermodynamical viewpoint." "Elementary complexions" spond to the "discrete configurations. Entropy = k log D where k is the atomic disorder. Entropy Physicists know that every isolated system changes of rest. being a measure of disorder. is The between entropy and probability given by the Boltzmann- Planck formula: Swhere S of is k In P the entropy. The entropic a measure of a difference is The value of entropy between an initial and a given by the formula: final state. But during reproduction two organisms have been pro- duced out of one. to a maximum Now probable relation entropy. The measure of entropy is radation energy. In a from a less probable to more state. in such is a a way state as to approach of a definite state It This a state of rest of equilibrium. and so does entropy. [89] . k the Boltzmann constant expressed in is. we have to consider the production or reproduction of order in its most general aspect. Order has increased.BIOLOGICAL ORDER AND ENTROPY Reproduction as well as maintenance involves a degradation of energy. that is. disorder. closed system evolving is related to probaa bility. corresponds not to loss but to a deg- change during such an irreversible process is the difference between the entropy at the end and the entropy at the beginning. As our subject is biological order. ergs per degree centigrade.

A living system cannot be closed. as a result of reproducentropy has decreased. How can we measure the atomic order. the negative entropy in a living organism? [90] . This negative entropy. And we may D is a a measure = k log D. If entropy S /flogl/D." represents the quality or grade if admit with Schrodinger that measure of atomic disorder. its reciprocal \/D will be of atomic order. the decrease of entropy corresponding to the increase of order.BIOLOGICAL ORDER Thus. the degree of atomic order or the availability for work. As already stated. should be clear that the live the organism plus a reasonable — system under consideration is that is. we have to measure corresponds to tion. nucleic bases. Has the over-all entropy of the system represented in the equation for metabolism and reproduction increased or decreased? To answer the question. Let us now consider the equation of reproduction: (Organism) 1 + food —> As (organism) 2 its + waste + heat The organism has synthesized building blocks — its essential metabolites. nor a fortiori isolated. negative entropy or negentropy N= Life and Entropy A An is system is said to be closed its when it does not carry on exchanges closed system is of matter with isolated surroundings. "negentropy. unlimited — supply of food. like the systems studied It by physicists. Thus. that is. A said to be when it does not interact in any its way with its surroundings. die. and amino acids — and has organized them into specific macromolecules. organism separated from surroundings will soon and therefore an abstraction. Physicists have found it convenient to have an expression for its reverse. living a result of the functioning of the machine. that is. Increase of order a decrease of entropy. of energy. entropy is a measure of atomic disorder. work has been produced and energy has been degraded. order has increased.

BIOLOGICAL ORDER AND ENTROPY Information and Negentropy This brings us to the negentropy principle of information. information is measured in entropy units: /i = 1. of any specific organization can thus be expressed For example. The information / is /i = K In Po based on scarcity. The /o = with Pq probable or possible outcomes. In 1929. This definition of information is The lower the probability. the higher the scarcity and the higher the information. And the demonstration has been provided that information corresponds to negative entropy. When number of possible answers when no some information is gained. the negentropy of a telephone net- work with 108 subscribers can be calculated to be 4 X 10""^. The similarity between the two formulas of entropy S = k\nP and information /i is = K In Po is obvious. that is to say. If the Boltzmann constant used. then is number of possible answers reduced.38 X 10-i«lnPo The information in entropy units. Leo Szilard discovered the existence of a connection between information and entropy. Information initial situation is a function of the ratio of possible answers before and after. One considers a problem involving information the tion" is a certain available. is What information? We shall follow Brillouin's analysis. [91] . one single outcome selected. The final situation is with Pi = 1. and "complete informais means only one possible answer.

organized in number a given sequence. we can calculate the of possible arrangements of the 10" units. This organization should perhaps be considered also in the calculation of the amount of information. or a nucleic acid. If we apply the formula /= it 1. But the problem of [92] . It contains a few thousand enzymes and a large number of other specific macromolecules. visualize the structural is a protein. Since this is so.BIOLOGICAL ORDER What This is. is the One could consider that the negentropy of an organism sum of the negentropy of its specific macromolecules. the negentropy. and that today there is only one. In the bacterium Escherichia coli. we have assumed that at the origin there were 4^0" possible outcomes. But an organism is much more than its genetic material. But organization of in all the problem can be simplified. terial is composed of 10" nucleic bases belonging to 4 categories which is the hereditary mamaterial. is the information. that is.38 X X 10-16 InPo follows that /= 1. The origin the macromolecules of an organism has its the organization of the hereditary nucleic acid. selected from among a large number of probabilities. of an organism? as pointed out by Brillouin in his classical book. explored field The Negentropy Physicists of Biological Organization know that in a machine the structural negentropy represents the information or organization of the machine. that is. ergs/°C. an unof research. This figure was obtained by considering only the genetic material. It is 4^^'. How can cell? we negentropy of an organism or of a Each species of macromolecule.38 10-16 In 410' The structural negentropy of the bacterium Escherichia coli is around 2 X 10 -^ entropy units. For the calculation of the information of our bacterium. part of the organization of the living machine. whether it is a polysaccharide.

radical development and a broad extension of the notion of entropy and to something quite different from the classical physical concept. its information will be destroyed. that this cannot be measured in terms of entropy units. functional coupling between the active units of a into account in the calculation of And Linschitz has proposed that this type of organization should also be taken what he calls a "biological I negenlike to tropy. one given molecule of guanine in a given gene is replaced by a molecule of adenine. This corresponds to a improbable system but a fitted for certain functions. one nucleic base of the genetic material sometimes replaced by another. The study cell. of the functional order has revealed a very high degree of nonstructural. original. for example. This alteration may cause the death of the organism.BIOLOGICAL ORDER AND ENTROPY physical entropy is even more complicated. the structural negentropy of the system. Nucleic acid therefore contains the blueprint for the [93] . The biologist feels that the functional order It is is an essential part of functional order is the living system. The value of the negentropy obviously does not give a measure of the efficiency of the system. And happens. would point by an example. As is a result of an accident. is the same. even in if the mutation is negentropy has remained the same. the negentropy of fected cell will die. and meaningless it from as will a purely thermodynamical point of view. The living organism is not only an Low system proposed Brillouin has probability and value are not synonymous. however. noninfected cell. It has ceased to be nothing has changed: the content consequence of the introduction' of the genetic material a cell/virus system is greater than the negentropy of the normal. a generalized negentropy principle that would take into account the value of the organization or information. As a of a virus. the altered organism is now unable to function normally and to reproduce. But the inalive. clear. the information. If. that is. than for the physicist. lethal. that the term "information" has for the biologist a different sense Remarks on "Genetic Information" The synthesis of a given specific protein is under the control of nucleic acid. be seen next." This illustrate this is certainly a very pertinent suggestion. For the physicist. But the mutation being lethal.

in the letters is know provided the number of contain the same is. that negative entropy. privileged. for each gene controls the synthesis of one given specific protein. Physicists account or a that the value of a message is not taken into measure of the information. the genes. The concept information. of the action of ultraviolet light on [94] . in a sense. But the biologist knows different from every other. contains information that negative entropy. "genetic information" refers to a given actual structure or order of the hereditary material and not to the negative entropy of this structure. included in the biological concept of genetic The gene is as a machine. and our position therefore. In an organism. The biologist currently speaks of information for the synthesis of a given enzyme. is have the same information. were organic molecules. or organization. that is. provided they the number of same content. The called genetic information. and the idea of quality. is the specific value.BIOLOGICAL ORDER orderly assemblage of amino acids into proteins. for evolution must be life also taken into account. for the biologist. Entropy and Evolution When order is functional order is considered. of probability has disappeared. But there is something more. for very biologists say that at the beginning there These were formed as a result good reasons. This should apply to all genetic information. the organization represents information. A theorem by Einstein random assemblage of letters both contain the same information. the same. It is believed that originated some 3 X 10^ years ago and that the organisms existing today are the consequence of a long evolution. But. and this is most regrettable. the hereditary organization has every right to be called information. In a machine. Yet. For the physicist. or structural negative entropy. Nothing is. Both the physicist and the biologist should be well aware of this situation. which that each gene is bases. positive is known about the origin of life. it appears that the biological considerably greater than the figure calculated by consider- ing only the probability of the base sequence in the hereditary material. for the production with blueprint is a probability 1 of a given specific structure.

As a result. producing new to certainly very different and better organisms that outgrew the original prototype. despite the fact that and a it contains and reproduces structural negative entropv^ very remarkable functional order. The Source In his remarkable of Orderliness everything that is book What Is Life.BIOLOGICAL ORDER AND ENTROPY simple inorganic compounds. The living system considered on a world-^\•ide scale cannot contradict the second principle. which today constitute the livincr world. and an organic gel endowed with catalytic properties eventually appeared. has the over-all entropy decreased The system organism is plus food might be visualized as as system but certainly not an isolated system. has doubled expense of an increase Is of entropy S. The first organism in turn started performing experiments. The it present biological order includes the historical experience its of the organism acquired during includes evolution that it evolution. and as a result the entropy increased. have been performed. at the same time structural and functional. a large number of species were selected. A large number of experiments. the catalytic "metabolizing" gel gave rise something able to reproduce itself. The organic molecules then assembled on some substrate such as colloidal clay. x'Vnd it is because is so highly improbable. As a result of a large number of experiments. This was the first organism. They necessarily correspond to an increase of entropy. This increase should be taken into account in any calculation of the negative entropy of the organism. When an organism metabolizes. Schrodinger states that going on in nature means an increase of entropy [95] . heat of the surrounding world is excreted. or increased? a closed AiV smaller or larger than AS? In other A\ords. due to the production of two at the organisms out of one. due to the degradation of energy. Let us consider again the equation ( Organism )i structural + food — >-( organism )2 + waste + heat The negentropy N. from any current living being. The organization of the actually living systems has been acquired at the expense of an increase of entropy. Everything unfit has been eliminated.

or orderliness. the source of orderliness is reasoning has been endorsed by Brillouin. The the real fuel for the maintenance of a biological point of A is steam engine is combustion of coal is the source of I movement of the machine. which is death. according to Schrodinger. according to Schrodinger." conserved and never gets food does not really matter. know that the formula "the organism feeds on negative entropy" has been applied to the maintenance of the living system. or organization. kept working by the combustion of coal. The organism can remain alive. it is only for the negentropy which losses from ical it and which is needed to make up for the work done. produces positive entropy. maintains its its order by "sucking orderliness" from environment. Schrodinger himself has is. Because biological order. since energy is lost. The only source of biological order maintenance as well as reproduction of biological order both depend on the same process. biological order. terms of negentropy. The organism. pointed out very pertinently that the characteristic of life is the prois duction of order from order." that is. It has been known for a long time that only an organism can give rise to an organism. or simple degradation processes in the living system. metabolism. reproduction being the other. For the animal. who writes. says Schrodinger. Negative entropy is. the the sunlight. orderliness [96] . This type of sources of orderliness are more-or-less complex organic compounds. that is. for the plant. but negentropy is the important The statements concerning negative entropy. a measure of order. just like information.BIOLOGICAL ORDER and that the living being continuously increases its entropy. "something very positive. namely. it not the source of the structure. can be expressed in Now. only by continuously drawing negative entropy from its environment. in Energy contained matter. thus approaching the state of maximum entropy. which is correlative to an increase of entropy. This is certainly all right." even when it maintenance. as life should be examined from view and also from a physical point of view. such statements alive by sucking applies only to as "The organism remains from its environment. But maintenance of the organism — that maintenance of biological order — is only one aspect of life. "If a living it can get due to mechan- organism needs food. might be misleading.

When light meets atoms. it has to perform oxidoreductions. In the original chemical bond. But it is clear that what is important in the molecule of sugar is not the structural negentropv of its atoms. one part work. Academic Press. Science and Inj orfnatio7i Theory. source The same seems to be true for the energy of the chemical bond. if it The as the steam engine does. [97] . Unless the energy of a chemical bond of food is utilized. as positive as negative entropy be. does not handle concepts of grade or logarithms of The organism energy of physicists I light or of handles atoms or molecules and the chemical bonds. L. New York. What is important is the energy of the chemicar bonds. have consulted decided that Schrodinger's formula was it did not make A general agreement was reached only on one point. as a result of a chemical reaction. the probability that these atoms are arranged in a certain order. and the other is potential entropy. whereas others claimed that sense at If all./ degraded. REFERENCES Brillouin. The organism probabilities.BIOLOGICAL ORDER AND ENTROPY wants to work. But. according to some physicists I have consulted. Negentropy is a grade of energy. such as the energy of light or of a chemical bond. But energy of high grade. just It might burn sugar. (1956). The energy of light allows the plant to separate hydrogen ions from the molecule of water. work is produced. for example. cannot be subdivided into positive and negative is available for entropy. When. some of the perfectly acceptable. It separates and binds atoms and molecules. Orderliness is a probability. Work. has to burn something. one cannot speak of free energy and of its entropic component. is produced at the expense of the energy of the chemical bonds. heat is pro- duced and a part of the energy degraded. as its it does not seem sound to state that the plant that uses light of energy feeds on the negative entropy of light. The organism is unable to make use of energy abstracto. with a correlative degradation of energy. /". Nevertheless. work is produced and energy is organism. The organism takes in food and burns it. fed with pellets of negative entropy. might even a physicist would succumb.

pp. Quastler. 251262. The information content of a bacterial cell. New York. Linschitz. (1953). Prigogine. H. (1958). University of Illinois Press. (1957). So7ne Principles of Energetics in Biochemical Reactions. [98] . 111. In Essays on the Use of Infor?/iation Theory in Biology. (1944)'. Paris. Cambridge. Wiener. M. Cybernetics. Hermann. University Press. I. edited by H. Introductioji to Springfield. E. I. What Is Life? The Physical Aspect of the Living Cell. Thernwdyna?nics of Irreversible Processes.BIOLOGICAL ORDER Klotz. Charles C Thomas. N. Urbana. Schrodinger. Academic Press. (1955).

so as their synthesis. a sequence of a few We have learned that nuckic acid. is a complex molecular society in which macromolecules and groups of macromolecules are interacting. synthesis of essential metabolites.CONCLUSION We small molecules. anism has developed. the stable. The an elaborate system of a feedback mechfunctioning of the organism reflects the [99] . the one cannot be produced The organism. is controlled by the product of their activity. The functioning of each enzyme or group of enzymes. are which of in turn the catalysts for the are. or better by the needs of the organism. or the cell. But. have learned that the very basis of is biological order. nucleic acid is the varvintT structure responsible for mutations and evolution. considered on the time scale of the world. Stability and variability reside in the is function of the genetic material hfe. This dual one of the important aspects of Nucleic acid is the blueprint for the synthesis of specific proteins. Each group possesses its system. own regulatory As a result of evolution. Nucleic acids and enzymes from in a functional point sense that view. same structure. of specificity and diversity. complementary macromolecules ^^'ithout the other. considered on is the time scale of the organism. All the groups and all the various regulating systems interact. permanent structure responsible for the specificity and for the reproduction of specificity.

a molecular tendency toward autonomy. enzymes. extracted from the cell — any structure. as a human being. Life can only be the appanage of the organism alive. I have In a acting. These viral is functions may be submitted to specific viral repressors. structural The genes of the host cell The pathological order the normal order is. and at the same time as a dependent part of a community. or the apparently consequence of toward an escape from cellular control. Waves and complementary aspects of the atom. Structures and functions are the complementary aspects of the organism. has to visualize himself as a person. Only organisms are The upsurge viral. structural genes. The study viral of viruses has revealed the existence of sequential controlling the functions viral genetic material autonomous reproduction of the and the synthesis of viral proteins. The genetic material of a virus sometimes under the control of the regulating mechanisms of the host. * * complex organized system.BIOLOGICAL ORDER high degree of coupling between regulating genes. which all macromolecules are interdependent The living organism is an integrated system of macromolecular its kind. Any [100] . Separated from its context — that is. just as may undergo structural recombination and exhibit functional interchanges.T. are sometimes under the control of the virus. as part of a whole. During three weeks. Normal and pathological entities types of dual order. whether cellular.I. The story of the cell/virus system is the story of the quadruple interactions between two different dual. operator genes. as an independent unit. cell is and metabolism. is just an organic molecule. repressors. is and functional. I had the privilege and pleasure to be part of M. is of molecular diseases. either a nucleic acid or a protein. subviral. in units. the units are necessarily interone of us. Such structures and functions able to reproduce particles are the a thing as living substance or living matter does not as a exist. The normal a structural and functional system of order. whole.

And I should like to express my admiration for this institute. disease. [101] . M.I.I. some powerful and efficient repressor-operator system. is But so far as can see.T. is perfectly healthy. an integrated system of strucI As I am a physician. the de- velopment of which has been so beautifully conceived by Karl Taylor Compton. This probably because the problem of the induction and repression is solved by some elaborate feedback mechanism. just like a living being. which no macromolecule can resist. I \ hich I already is somehow and a as suspected. namely. am interested in have tried to detect some sign of I pathological process. that M.CONCLUSION discovered something tures and functions..T.

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